Evaluation of hemispheric dominance for language using functional MRI: A comparison with positron emission tomography

The utility of a conventional (i.e., nonecho-planar) functional magnetic resonance imaging (fMRI) technique to determine hemispheric dominance for language was assessed using a semantic generation task in which subjects were presented with a series of nouns and generated aloud a verb for each one. A direct comparison of the fMRI results with positron emission tomography (PET), acquired from the same subjects, was also performed. When analyzed by group averaging, the results of this work were in concordance with those of previous PET studies, showing a left hemispheric dominance for language. Analyzed on an individual basis, 7 out of 9 subjects were left-hemisphere dominant and 2 subjects were right-hemisphere dominant; this applied with both PET and fMRI methods. The direct comparison between PET and fMRI further demonstrated that fMRI can replicate PET findings. Of the total activation foci detected by PET, 92% were observed by fMRI. On the other hand, fMRI reported 64% more activations than did PET. This may reflect differences in procedure, spatial resolution, sensitivity, and the underlying physiological mechanism between PET and fMRI. This study suggests that the fMRI technique, even using conventional MRI scanners, could be clinically useful in evaluating hemispheric dominance for language on an individual subject-by-subject basis. Hum. Brain Mapping 6:42–58, 1998. © 1998 Wiley-Liss, Inc.     

Regional differences in the developmental trajectory of lateralization of the language network

The timing and developmental factors underlying the establishment of language dominance are poorly understood. We investigated the degree of lateralization of traditional frontotemporal and modulatory prefrontal‐cerebellar regions of the distributed language network in children (n = 57) ages 4 to 12—a critical period for language consolidation. We examined the relationship between the strength of language lateralization and neuropsychological measures and task performance. The fundamental language network is established by four with ongoing maturation of language functions as evidenced by strengthening of lateralization in the traditional frontotemporal language regions; temporal regions were strongly and consistently lateralized by age seven, while frontal regions had greater variability and were less strongly lateralized through age 10. In contrast, the modulatory prefrontal‐cerebellar regions were the least strongly lateralized and degree of lateralization was not associated with age. Stronger core language skills were significantly correlated with greater right lateralization in the cerebellum. Hum Brain Mapp 35:270–284, 2014. © 2012 Wiley Periodicals, Inc.     

Working memory representation in atypical language dominance

One of the most important factors controlling material specific processing in the human brain is language dominance, i.e. hemispheric specialization in semantic processes. Although previous studies have shown that lateralized long-term memory processes in the medial temporal lobes are modified in subjects with atypical (right) language dominance, the effect of language dominance on the neural basis of working memory (WM) has remained unknown. Here, we used functional MRI (fMRI) to study the impact of language dominance on the neural representation of WM. We conducted an n-back task in three different load conditions and with both verbal and nonverbal (spatial) material in matched groups of left and right language dominant subjects. This approach allowed us to investigate regions showing significant interactions between language dominance and material. Overall, right dominant subjects showed an increased inter-individual variability of WM-related activations. Verbal WM involved more pronounced activation of the left fusiform cortex in left dominant subjects and of the right inferior parietal lobule in the right dominant group. Spatial WM, on the other hand, induced activation of right hemispheric regions in left dominant subjects, but no specific activations in right dominant subjects. Taken together, these findings indicate that the neural basis of verbal WM processes depends on language dominance and is more mutable in right dominant subjects. The increased variability in right dominant subjects strongly suggests that a standard network of material-dependent WM processes exists in left dominant subjects, and that right dominant subjects use variable alternative networks.     

fMRI study of language lateralization in children and adults

Language lateralization in the brain is dependent on family history of handedness, personal handedness, pathology, and other factors. The influence of age on language lateralization is not completely understood. Increasing left lateralization of language with age has been observed in children, while the reverse has been noted in healthy young adults. It is not known whether the trend of decreasing language lateralization with age continues in the late decades of life and at what age the inflection in language lateralization trend as a function of age occurs. In this study, we examined the effect of age on language lateralization in 170 healthy right-handed children and adults ages 5-67 using functional MRI (fMRI) and a verb generation task. Our findings indicate that language lateralization to the dominant hemisphere increases between the ages 5 and 20 years, plateaus between 20 and 25 years, and slowly decreases between 25 and 70 years.     

Sensory‐motor brain network connectivity for speech comprehension

The act of listening to speech activates a large network of brain areas. In the present work, a novel data‐driven technique (the combination of independent component analysis and Granger causality) was used to extract brain network dynamics from an fMRI study of passive listening to Words, Pseudo‐Words, and Reverse‐played words. Using this method we show the functional connectivity modulations among classical language regions (Broca's and Wernicke's areas) and inferior parietal, somatosensory, and motor areas and right cerebellum. Word listening elicited a compact pattern of connectivity within a parieto‐somato‐motor network and between the superior temporal and inferior frontal gyri. Pseudo‐Word stimuli induced activities similar to the Word condition, which were characterized by a highly recurrent connectivity pattern, mostly driven by the temporal lobe activity. Also the Reversed‐Word condition revealed an important influence of temporal cortices, but no integrated activity of the parieto‐somato‐motor network. In parallel, the right cerebellum lost its functional connection with motor areas, present in both Word and Pseudo‐Word listening. The inability of the participant to produce the Reversed‐Word stimuli also evidenced two separate networks: the first was driven by frontal areas and the right cerebellum toward somatosensory cortices; the second was triggered by temporal and parietal sites towards motor areas. Summing up, our results suggest that semantic content modulates the general compactness of network dynamics as well as the balance between frontal and temporal language areas in driving those dynamics. The degree of reproducibility of auditory speech material modulates the connectivity pattern within and toward somatosensory and motor areas. Hum Brain Mapp, 2010. © 2009 Wiley‐Liss, Inc.     

Determination of crossed language dominance: dissociation of language lateralization within the temporoparietal cortex

‘Crossed language dominance’ is a rare form of language lateralization, characterized by a dissociation of anterior and posterior language regions. We present the case of a healthy subject whose language lateralization pattern, as assessed by functional magnetic resonance imaging, is reliably characterized as crossed language dominance based on a word generation task, but typical left-lateralized when a semantic decision task is applied. A single language task is therefore not sufficient to characterize language lateralization, at least not for subjects with rare forms of language dominance. In the pre-surgical diagnostic of language lateralization, several language tasks tapping into different aspects of language functions should be applied.     

Presurgical functional transcranial Doppler sonography (fTCD) with intravenous echo enhancing agent SonoVue enables determination of language lateralization in epilepsy patients with poor temporal bone windows

Presurgical determination of language lateralization is important for planning and outcome estimation of neurosurgical interventions in patients with drug-refractory epilepsy. Functional transcranial Doppler sonography (fTCD) provides an established measure for language lateralization using the temporal bone windows for continuous recording of the cerebral blood flow velocity (CBFV) in both middle cerebral arteries (MCAs). However, because of insufficient temporal bone windows, fTCD cannot be applied properly in every patient. Here, we established stable and sufficient CBFV signals in both MCAs using continuous intravenous application of echo-enhancing agent SonoVue in 7 of 10 patients with poor temporal bone windows and were thus able to determine language lateralization. We conclude that the application of SonoVue can solve one principal disadvantage of fTCD and improves the applicability of the technique as a presurgical functional language lateralization procedure.     

Arcuate fasciculus asymmetry has a hand in language function but not handedness

The importance of relationships between handedness, language lateralization and localization, and white matter tracts for language performance is unclear. The goal of the study was to investigate these relationships by examining arcuate fasciculus (AF) structural asymmetry (DTI) and functional asymmetry (fMRI) in language circuits, handedness, and linguistic performance. A large sample of right‐handed (n = 158) and atypical‐handed (n = 82) healthy adults underwent DTI at 3 T to assess number of streamlines and fractional anisotropy (FA) of the AF, and language fMRI. Language functions were assessed using standard tests of vocabulary, naming, verbal fluency, and complex ideation. Laterality indices (LIs) illustrated degree of asymmetry and lateralization patterns for the AF (streamlines and FA) and verb generation fMRI. Both handedness groups showed leftward lateralization bias for streamline and fMRI LIs and symmetry for FA LI. The proportion of subjects with left, right, or symmetric lateralization were similar between groups if based on AF LIs, but differed if based on fMRI LIs (p = 0.0016). Degree of right‐handedness was not associated with AF lateralization, but was associated with fMRI language lateralization (p = 0.0014). FA LI was not associated with performance on language assessments, but streamline LI was associated with better vocabulary and complex ideation performance in atypical‐handed subjects (p = 0.022 and p = 0.0098, respectively), and better semantic fluency in right‐handed subjects (p = 0.047); however, these did not survive multiple comparisons correction. We provide evidence that AF asymmetry is independent of hand preference, and while degree of right‐handedness is associated with hemispheric language lateralization, the majority of atypical‐handed individuals are left‐lateralized for language. Hum Brain Mapp 37:3297–3309, 2016. © 2016 Wiley Periodicals, Inc .     

Sensorimotor,language, and working memory representation within the human cerebellum

The cerebellum is involved in a wide range of behaviours. A key organisational principle from animal studies is that somatotopically corresponding sensory input and motor output reside in the same cerebellar cortical areas. However, compelling evidence for a similar arrangement in humans and whether it extends to cognitive functions is lacking. To address this, we applied cerebellar optimised whole‐brain functional MRI in 20 healthy subjects. To assess spatial overlap within the sensorimotor and cognitive domains, we recorded activity to a sensory stimulus (vibrotactile) and a motor task; the Sternberg verbal working memory (VWM) task; and a verb generation paradigm. Consistent with animal data, sensory and motor activity overlapped with a somatotopic arrangement in ipsilateral areas of the anterior and posterior cerebellum. During the maintenance phase of the Sternberg task, a positive linear relationship between VWM load and activity was observed in right Lobule VI, extending into Crus I bilaterally. Articulatory movement gave rise to bilateral activity in medial Lobule VI. A conjunction of two independent language tasks localised activity during verb generation in right Lobule VI‐Crus I, which overlapped with activity during VWM. These results demonstrate spatial compartmentalisation of sensorimotor and cognitive function in the human cerebellum, with each area involved in more than one aspect of a given behaviour, consistent with an integrative function. Sensorimotor localisation was uniform across individuals, but the representation of cognitive tasks was more variable, highlighting the importance of individual scans for mapping higher order functions within the cerebellum.     

Left‐handed musicians show a higher probability of atypical cerebral dominance for language

Music processing and right hemispheric language lateralization share a common network in the right auditory cortex and its frontal connections. Given that the development of hemispheric language dominance takes place over several years, this study tested whether musicianship could increase the probability of observing right language dominance in left‐handers. Using a classic fMRI language paradigm, results showed that atypical lateralization was more predominant in musicians (40%) than in nonmusicians (5%). Comparison of left‐handers with typical left and atypical right lateralization revealed that: (a) atypical cases presented a thicker right pars triangularis and more gyrified left Heschl's gyrus; and (b) the right pars triangularis of atypical cases showed a stronger intra‐hemispheric functional connectivity with the right angular gyrus, but a weaker interhemispheric functional connectivity with part of the left Broca's area. Thus, musicianship is the first known factor related to a higher prevalence of atypical language dominance in healthy left‐handed individuals. We suggest that differences in the frontal and temporal cortex might act as shared predisposing factors to both musicianship and atypical language lateralization.     

Cerebellar activation and lateralization during verbal and visuospatial tasks as revealed by fMRI

BACKGROUND: Many studies concerning cerebral activation and lateralization of cognitive functions are being conducted. Cerebellar function has been much researched with reference to high-level cognitive processing, but has been barely researched in systematization and diversification. In particular, cerebellar lateralization has never been researched, in comparison to cerebral lateralization for which reliable results have been reported. OBJECTIVE: This study ascertains cerebellar activation and its lateralization in relation to verbal and visuospatial tasks, using functional magnetic resonance imaging (fMRI). DESIGN, TIME, AND SETTING: A block design for functional Magnetic Resonance Imaging (fMRI) observation. This study was performed at the fMRI Laboratory, Brain Science Research Center, Korea Advanced Institute of Science and Technology from May 2006 to September 2008. PARTICIPANTS: Sixteen healthy male college students (23.3 ± 0.5 years) and another sixteen healthy male college students (21.5 ± 2.3 years) participated in this fMRI study of verbal and visuospatial tasks, respectively. METHODS: The verbal and visuospatial tasks were presented while functional brain images were acquired using a 3T fMRI system (ISOL Technology, Korea). The verbal analogy testing required the subject to select the word which had the same relationship as one of the given words. The verbal antonym testing required the subject to select the word which had a different meaning among 4 words. The visuospatial tasks involved selecting a shape that corresponded to a given figure using four examples, as well as selecting a development figure of a diagram. MAIN OUTCOME MEASURE: The double subtraction method was used to analyze the differences of the cerebellar activation between the two cognition tasks. The numbers of activated voxels were calculated in bilateral cerebellums for the two tasks. The lateralization index of the cerebellum was calculated for each task. RESULTS: The bilateral hemisphere lobule VI and IX, the right hemisphere lobule VIII, the bilateral hemisphere lobule crus I, and the vermis lobule IV, V and VI are closely related to verbal tasks in comparison to visuospatial tasks. Conversely, the bilateral hemisphere lobule IV and V, and the right hemisphere lobule VI were closely related to visuospatial tasks compared to verbal tasks. There was no great difference between the bilateral cerebellums in the numbers of activated voxels for the tasks and cerebellar lateralization was not observed. CONCLUSION: In the case of the cerebellum, the activation region was different between the verbal and visuospatial tasks, but lateralization was not different.     

Hemispheric specialization for language in children with different types of specific language impairment

The aim of the study was to investigate whether children with Specific Language Impairment (SLI) show reduced left hemisphere specialization for language and, if so, whether it is associated with a deficit in phonological encoding and a specific type of SLI (Mixed Receptive-Expressive, Expressive, Phonological). We adopted two dichotic listening paradigms, which differed in the phonological similarity of the presented words (Fused and Non-Fused dichotic words tests), as well as a phonological working memory test. Participants included 34 pre-school and school age children affected by SLI. On the dichotic tests, as a group the children with SLI showed a reduced pattern of left hemisphere specialization for language compared to age-matched normal children, with significant differences only in the Fused condition. However, the pattern of hemispheric specialization varied depending on the type of SLI, with reduced left hemisphere specialization in the Expressive type and, to a lesser extent, in the Phonological type of SLI, but not in the Receptive-Expressive type. The three subgroups also differed in phonological processing abilities and the incidence of a positive family history for language disorders: the Receptive-Expressive group performed worse on the working memory and dichotic tests and the Expressive and Phonological groups presented high frequency for familial language disorder. These results suggest that different subtypes of SLI are not different manifestations of the same underlying disorder, but represent pathological conditions that have distinct markers both at the behavioral and neurofunctional level.     

Neuropsychological and functional MRI studies provide converging evidence of anterior language dysfunction in BECTS

Purpose:   Benign childhood epilepsy with centrotemporal spikes (BECTS) is the most common epilepsy syndrome of childhood and can be associated with language difficulties. The exact profile of these difficulties and their neurofunctional underpinnings, however, are not yet clear.
Methods:   To further understand the impact of the BECTS syndrome on language, we assessed language performance using standard neuropsychological measures, and patterns of language lateralization using functional magnetic resonance imaging (fMRI) in children with typical BECTS (n = 20) and healthy controls (n = 20).
Results:   The fMRI analyses revealed that language-related activation was less lateralized to the left hemisphere in anterior brain regions in the patients relative to the control group. This finding was consistent with decreased performance in the BECTS group compared to the control group on the neuropsychological measure most dependent on the integrity of anterior aspects of the language axis, namely, sentence production.
Discussion:   The converging lines of evidence from the neuropsychological and activation methodologies support the view that BECTS is associated with language difficulties that are regional, and anterior, in nature.     

Cerebral mechanisms of prosodic sensory integration using low-frequency bands of connected speech

Even if speech perception has been reported to involve both left and right hemispheres, converging data have posited the existence of a functional asymmetry at the level of secondary auditory cortices. Using fMRI in 12 right-handed French men listening passively to long connected speech stimuli, we addressed the question of neuronal networks involved in the integration of low frequency bands of speech by comparing 1) differences in brain activity in two listening conditions (FN, NF) differing in the integration of pitch modulations (in FN, low frequencies, obtained by a low-pass filter, are addressed to the left ear while the whole acoustic message is simultaneously addressed to the right ear, NF being the reverse position); 2) differences in brain activity induced by high and low degrees of prosodic expression (expressive vs. flat); and 3) effects of the same connected speech stimulus in the two listening conditions. Each stimulus induced a specific cerebral network, the flat one weakening activations which were mainly reduced to the bilateral STG for both listening conditions. In the expressive condition, the specific sensory integration FN results in an increase of the articulatory loop and new recruitments such as right BA6-44, left BA39-40, the left posterior insula and the bilateral BA30. This finding may be accounted for by the existence of temporal windows differing both in length and in acoustic cues decoding, strengthening the "asymmetric sampling in time" hypothesis posited by Poeppel (Speech Commun 2003; 41:245-255). Such an improvement of prosodic integration could find applications in the rehabilitation of some speech disturbances.     

The signer and the sign: Cortical correlates of person identity and language processing from point-light displays

In this study, the first to explore the cortical correlates of signed language (SL) processing under point-light display conditions, the observer identified either a signer or a lexical sign from a display in which different signers were seen producing a number of different individual signs. Many of the regions activated by point-light under these conditions replicated those previously reported for full-image displays, including regions within the inferior temporal cortex that are specialised for face and body-part identification, although such body parts were invisible in the display. Right frontal regions were also recruited – a pattern not usually seen in full-image SL processing. This activation may reflect the recruitment of information about person identity from the reduced display. A direct comparison of identify-signer and identify-sign conditions showed these tasks relied to a different extent on the posterior inferior regions. Signer identification elicited greater activation than sign identification in (bilateral) inferior temporal gyri (BA 37/19), fusiform gyri (BA 37), middle and posterior portions of the middle temporal gyri (BAs 37 and 19), and superior temporal gyri (BA 22 and 42). Right inferior frontal cortex was a further focus of differential activation (signer > sign).These findings suggest that the neural systems supporting point-light displays for the processing of SL rely on a cortical network including areas of the inferior temporal cortex specialized for face and body identification. While this might be predicted from other studies of whole body point-light actions (Vaina, Solomon, Chowdhury, Sinha, & Belliveau, 2001) it is not predicted from the perspective of spoken language processing, where voice characteristics and speech content recruit distinct cortical regions (Stevens, 2004) in addition to a common network. In this respect, our findings contrast with studies of voice/speech recognition (Von Kriegstein, Kleinschmidt, Sterzer, & Giraud, 2005). Inferior temporal regions associated with the visual recognition of a person appear to be required during SL processing, for both carrier and content information.     

Speech listening specifically modulates the excitability of tongue muscles: a TMS study

The precise neural mechanisms underlying speech perception are still to a large extent unknown. The most accepted view is that speech perception depends on auditory-cognitive mechanisms specifically devoted to the analysis of speech sounds. An alternative view is that, crucial for speech perception, it is the activation of the articulatory (motor) gestures that generate these sounds. The listener understands the speaker when his/her articulatory gestures are activated (motor theory of speech perception). Here, by using transcranial magnetic stimulation (TMS), we demonstrate that, during speech listening, there is an increase of motor-evoked potentials recorded from the listeners' tongue muscles when the presented words strongly involve, when pronounced, tongue movements. Although these data do not prove the motor theory of speech perception, they demonstrate for the first time that word listening produces a phoneme specific activation of speech motor centres.     

Presurgical language fMRI in patients with drug-resistant epilepsy: effects of task performance

PURPOSE: To determine whether language functional magnetic resonance imaging (fMRI) before epilepsy surgery can be similarly interpreted in patients with greatly different performance levels. METHODS: An fMRI paradigm using a semantic decision task with performance control and a perceptual control task was applied to 226 consecutive patients with drug-resistant localization-related epilepsy during their presurgical evaluations. The volume of activation and lateralization in an inferior frontal and a temporoparietal area was assessed in correlation with individual performance levels. RESULTS: We observed differential effects of task performance on the volume of activation in the inferior frontal and the temporoparietal region of interest, but performance measures did not correlate with the lateralization of activation. CONCLUSIONS: fMRI, as applied here, in patients with a wide range of cognitive abilities, can be interpreted regarding language lateralization in a similar way.     

Emotional language is all right: Emotional prosody reduces hemispheric asymmetry for linguistic processing

ABSTRACT

In an influential paper, Bryden and MacRae [(1989). Dichotic laterality effects obtained with emotional words. Neuropsychiatry, Neuropsychology, and Behavioral Neurology, 1, 171–176] introduced a dichotic listening task that allowed for the assessment of linguistic and prosodic processing asymmetries using the same stimuli. The task produces a robust right ear advantage (REA) for linguistic processing (identifying a target word), and a left ear advantage for emotional processing (identifying a target prosody). Here, we adapted this paradigm to determine whether and how the presence of emotional prosody might modulate hemispheric asymmetry for linguistic processing. Participants monitored for a target word among dichotic stimuli consisting of two different words, but spoken in the same emotional prosody—neutral, angry, happy, sad, or fearful. A strong REA was observed when the words were spoken in neutral prosody, which was attenuated for all the emotional prosodies. There were no differences in the ear advantage as a function of valence or discrete emotion, indicating that all emotions had similar effects. Findings are consistent with the hypothesis that the right hemisphere is better able to process speech when it carries emotional prosody. Implications for understanding of right hemisphere language functions are discussed.     

A voxel‐based asymmetry study of the relationship between hemispheric asymmetry and language dominance in Wada tested patients

Determining the anatomical basis of hemispheric language dominance (HLD) remains an important scientific endeavor. The Wada test remains the gold standard test for HLD and provides a unique opportunity to determine the relationship between HLD and hemispheric structural asymmetries on MRI. In this study, we applied a whole‐brain voxel‐based asymmetry (VBA) approach to determine the relationship between interhemispheric structural asymmetries and HLD in a large consecutive sample of Wada tested patients. Of 135 patients, 114 (84.4%) had left HLD, 10 (7.4%) right HLD, and 11 (8.2%) bilateral language representation. Fifty‐four controls were also studied. Right‐handed controls and right‐handed patients with left HLD had comparable structural brain asymmetries in cortical, subcortical, and cerebellar regions that have previously been documented in healthy people. However, these patients and controls differed in structural asymmetry of the mesial temporal lobe and a circumscribed region in the superior temporal gyrus, suggesting that only asymmetries of these regions were due to brain alterations caused by epilepsy. Additional comparisons between patients with left and right HLD, matched for type and location of epilepsy, revealed that structural asymmetries of insula, pars triangularis, inferior temporal gyrus, orbitofrontal cortex, ventral temporo‐occipital cortex, mesial somatosensory cortex, and mesial cerebellum were significantly associated with the side of HLD. Patients with right HLD and bilateral language representation were significantly less right‐handed. These results suggest that structural asymmetries of an insular‐fronto‐temporal network may be related to HLD.