Lise Van der Haegen

On the other hand: including left-handers in cognitive neuroscience and neurogenetics.

Left-handers are often excluded from study cohorts in neuroscience and neurogenetics in order to reduce variance in the data. However, recent investigations have shown that the inclusion or targeted recruitment of left-handers can be informative in studies on a range of topics, such as cerebral lateralization and the genetic underpinning of asymmetrical brain development. Left-handed individuals represent a substantial portion of the human population and therefore left-handedness falls within the normal range of human diversity; thus, it is important to account for this variation in our understanding of brain functioning. We call for neuroscientists and neurogeneticists to recognize the potential of studying this often-discarded group of research subjects.

Complementary hemispheric specialization for language production and visuospatial attention

Language production and spatial attention are the most salient lateralized cerebral functions, and their complementary specialization has been observed in the majority of the population. To investigate whether the complementary specialization has a causal origin (the lateralization of one function causes the opposite lateralization of the other) or rather is a statistical phenomenon (different functions lateralize independently), we determined the lateralization for spatial attention in a group of individuals with known atypical right hemispheric (RH) lateralization for speech production, based on a previous large-scale screening of left-handers. We show that all 13 participants with RH language dominance have left-hemispheric dominance for spatial attention, and all but one of 16 participants with left-hemispheric language dominance are RH dominant for spatial attention. Activity was observed in the dorsal fronto-parietal pathway of attention, including the inferior parietal sulcus and superior parietal lobule, the frontal eye-movement field, and the inferior frontal sulcus/gyrus, and these regions functionally colateralized in the hemisphere dominant for attention, independently of the side of lateralization. Our results clearly support the Causal hypothesis about the complementary specialization, and we speculate that it derives from a longstanding evolutionary origin. We also suggest that the conclusions about lateralization based on an unselected sample of the population and laterality assessment using coarse functional transcranial Doppler sonography should be interpreted with more caution.

A surface-based analysis of language lateralization and cortical asymmetry

Among brain functions, language is one of the most lateralized. Cortical language areas are also some of the most asymmetrical in the brain. An open question is whether the asymmetry in function is linked to the asymmetry in anatomy. To address this question, we measured anatomical asymmetry in 34 participants shown with fMRI to have language dominance of the left hemisphere (LLD) and 21 participants shown to have atypical right hemisphere dominance (RLD). All participants were healthy and left-handed, and most (80%) were female. Gray matter (GM) volume asymmetry was measured using an automated surface-based technique in both ROIs and exploratory analyses. In the ROI analysis, a significant difference between LLD and RLD was found in the insula. No differences were found in planum temporale (PT), pars opercularis (POp), pars triangularis (PTr), or Heschls gyrus (HG). The PT, POp, insula, and HG were all significantly left lateralized in both LLD and RLD participants. Both the positive and negative ROI findings replicate a previous study using manually labeled ROIs in a different cohort [Keller, S. S., Roberts, N., Garcia-Finana, M., Mohammadi, S., Ringelstein, E. B., Knecht, S., et al. Can the language-dominant hemisphere be predicted by brain anatomy? Journal of Cognitive Neuroscience, 23, 2013–2029, 2011]. The exploratory analysis was accomplished using a new surface-based registration that aligns cortical folding patterns across both subject and hemisphere. A small but significant cluster was found in the superior temporal gyrus that overlapped with the PT. A cluster was also found in the ventral occipitotemporal cortex corresponding to the visual word recognition area. The surface-based analysis also makes it possible to disentangle the effects of GM volume, thickness, and surface area while removing the effects of curvature. For both the ROI and exploratory analyses, the difference between LLD and RLD volume laterality was most strongly driven by differences in surface area and not cortical thickness. Overall, there were surprisingly few differences in GM volume asymmetry between LLD and RLD indicating that gross morphometric asymmetry is only subtly related to functional language laterality.

Praxis and language are linked: Evidence from co-lateralization in individuals with atypical language dominance

We determined the neural correlates of word generation and tool use pantomiming in healthy subjects with typical (n=10) or atypical (n=10) language dominance to investigate similarities in response pattern and hemispheric specialization between language and praxis. All typical language dominant volunteers also revealed left hemisphere changes during tool use pantomiming in prefrontal, premotor, and posterior parietal regions. All atypical language dominant participants displayed right hemisphere engagement for tool use. Co-lateralization of the language and praxis networks was observed on group and individual level, regardless of the participants handedness. Activation maps of the word generation and tool use pantomiming contrasts displayed overlap in five cortical regions: supplementary motor area, dorsal and ventral premotor cortex, dorsolateral prefrontal cortex, and posterior parietal cortex. Individual lateralization indices were calculated for each region and revealed significant positive group correlations between .51 and .95 with every other region within the paradigms. Positive cross-task correlations ranged between .72 (supplementary motor complex) and .97 (dorsal premotor cortex) and illustrate that the strength of hemispheric specialization of one task significantly predicts the side and degree of lateralization of the other task, suggesting a functional and topographic link between language and praxis. These findings support models that link gestures and speech to explain the evolution of human language. We argue that the existence of a common and co-lateralized network underlying the production of complex learned movement, whether it be speech or tool use, may represent the evolutionary remnant of a neural system out of which proto-sign and proto-speech co-evolved.

New human-specific brain landmark: the depth asymmetry of superior temporal sulcus

Significance In the human brain, from early in development through to adulthood, the superior temporal sulcus is deeper in the right than the left cerebral hemisphere in the area ventral of Heschl’s gyrus. Irrespective of gender, handedness, and language lateralization, and present in several pathologies, this asymmetry is widely shared among the human population. Its appearance early in life suggests strong genetic control over this part of the brain. In contrast, the asymmetry is barely visible in chimpanzees. Thus this asymmetry probably is a key locus to look for variations in gene expression among the primate lineage that have favored the evolution of crucial cognitive abilities sustained by this sulcus in our species, namely communication and social cognition. Identifying potentially unique features of the human cerebral cortex is a first step to understanding how evolution has shaped the brain in our species. By analyzing MR images obtained from 177 humans and 73 chimpanzees, we observed a human-specific asymmetry in the superior temporal sulcus at the heart of the communication regions and which we have named the “superior temporal asymmetrical pit” (STAP). This 45-mm-long segment ventral to Heschl’s gyrus is deeper in the right hemisphere than in the left in 95% of typical human subjects, from infanthood till adulthood, and is present, irrespective of handedness, language lateralization, and sex although it is greater in males than in females. The STAP also is seen in several groups of atypical subjects including persons with situs inversus, autistic spectrum disorder, Turner syndrome, and corpus callosum agenesis. It is explained in part by the larger number of sulcal interruptions in the left than in the right hemisphere. Its early presence in the infants of this study as well as in fetuses and premature infants suggests a strong genetic influence. Because this asymmetry is barely visible in chimpanzees, we recommend the STAP region during midgestation as an important phenotype to investigate asymmetrical variations of gene expression among the primate lineage. This genetic target may provide important insights regarding the evolution of the crucial cognitive abilities sustained by this sulcus in our species, namely communication and social cognition.

Further fMRI validation of the visual half field technique as an indicator of language laterality: A large-group analysis

The best established lateralized cerebral function is speech production, with the majority of the population having left hemisphere dominance. An important question is how to best assess the laterality of this function. Neuroimaging techniques such as functional Magnetic Resonance Imaging (fMRI) are increasingly used in clinical settings to replace the invasive Wada-test. We evaluated the usefulness of behavioral visual half field (VHF) tasks for screening a large sample of healthy left-handers. Laterality indices (LIs) calculated on the basis of the latencies in a word and picture naming VHF task were compared to the brain activity measured in a silent word generation task in fMRI (pars opercularis/BA44 and pars triangularis/BA45). Results confirmed the usefulness of the VHF-tasks as a screening device. None of the left-handed participants with clear right visual field (RVF) advantages in the picture and word naming task showed right hemisphere dominance in the scanner. In contrast, 16/20 participants with a left visual field (LVF) advantage in both word and picture naming turned out to have atypical right brain dominance. Results were less clear for participants who failed to show clear VHF asymmetries (below 20 ms RVF advantage and below 60 ms LVF advantage) or who had inconsistent asymmetries in picture and word naming. These results indicate that the behavioral tasks can mainly provide useful information about the direction of speech dominance when both VHF differences clearly point in the same direction.

Colateralization of Broca’s area and the visual word form area in left-handers: fMRI evidence

Language production has been found to be lateralized in the left hemisphere (LH) for 95% of right-handed people and about 75% of left-handers. The prevalence of atypical right hemispheric (RH) or bilateral lateralization for reading and colateralization of production with word reading laterality has never been tested in a large sample. In this study, we scanned 57 left-handers who had previously been identified as being clearly left (N=30), bilateral (N=7) or clearly right (N=20) dominant for speech on the basis of fMRI activity in the inferior frontal gyrus (pars opercularis/pars triangularis) during a silent word generation task. They were asked to perform a lexical decision task, in which words were contrasted against checkerboards, to test the lateralization of reading in the ventral occipitotemporal region. Lateralization indices for both tasks correlated significantly (r=0.59). The majority of subjects showed most activity during lexical decision in the hemisphere that was identified as their word production dominant hemisphere. However, more than half of the sample (N=31) had bilateral activity for the lexical decision task without a clear dominant role for either the LH or RH, and three showed a crossed frontotemporal lateralization pattern. These findings have consequences for neurobiological models relating phonological and orthographic processes, and for lateralization measurements for clinical purposes.

Blinking predicts enhanced cognitive control.

Recent models have suggested an important role for neuromodulation in explaining trial-to-trial adaptations in cognitive control. The adaptation-by-binding model (Verguts & Notebaert, Psychological review, 115(2), 518–525, 2008), for instance, suggests that increased cognitive control in response to conflict (e.g., incongruent flanker stimulus) is the result of stronger binding of stimulus, action, and context representations, mediated by neuromodulators like dopamine (DA) and/or norepinephrine (NE). We presented a flanker task and used the Gratton effect (smaller congruency effect following incongruent trials) as an index of cognitive control. We investigated the Gratton effect in relation to eye blinks (DA related) and pupil dilation (NE related). The results for pupil dilation were not unequivocal, but eye blinks clearly modulated the Gratton effect: The Gratton effect was enhanced after a blink trial, relative to after a no-blink trial, even when controlling for correlated variables. The latter suggests an important role for DA in cognitive control on a trial-to-trial basis.

Speech dominance is a better predictor of functional brain asymmetry than handedness: A combined fMRI word generation and behavioral dichotic listening study☆

An unresolved issue in behavioral studies of hemispheric asymmetry is why both left-handers and right-handers show a right ear advantage at the group level. In the present study we screened left-handers for left- versus right-hemisphere speech dominance with fMRI by comparing right versus left hemisphere frontal lobe activity (in Brocas area) in a silent word generation task. A left hemisphere dominant right-handed control group was included as well. All participants took part in a dichotic listening task with consonant-vowel syllables. The results showed that left-handers and right-handers with left-hemisphere speech dominance showed a right ear advantage. However, the left-handers with right hemisphere speech dominance had a left ear advantage. Thus, at the group level the direction of the ear advantage in dichotic listening was predicted by language dominance but not by hand preference. At the individual level, the dichotic task we used showed more variability than the fMRI results. Further research will have to indicate whether this is a feature inherent to dichotic listening, or whether the variability is due to alternative explanations such as a more bilateral representation of speech perception compared to speech production.

How does interhemispheric communication in visual word recognition work? Deciding between early and late integration accounts of the split fovea theory.

It has recently been shown that interhemispheric communication is needed for the processing of foveally presented words. In this study, we examine whether the integration of information happens at an early stage, before word recognition proper starts, or whether the integration is part of the recognition process itself. Two lexical decision experiments are reported in which words were presented at different fixation positions. In Experiment 1, a masked form priming task was used with primes that had two adjacent letters transposed. The results showed that although the fixation position had a substantial influence on the transposed letter priming effect, the priming was not smaller when the transposed letters were sent to different hemispheres than when they were projected to the same hemisphere. In Experiment 2, stimuli were presented that either had high frequency hemifield competitors or could be identified unambiguously on the basis of the information in one hemifield. Again, the lexical decision times did not vary as a function of hemifield competitors. These results are consistent with the early integration account, as presented in the SERIOL model of visual word recognition.