One of the oldest findings in human neuroscience is that the brain control of language is lateralized to the left hemisphere (Broca, 1861; Wernicke, 1874; Price, 2012). However, not all individuals present a typical organization of this function. Some left-handers (22–24%) show an atypical lateralization of language – right or ambilateral (Pujol et al., 1999; Szaflarski et al., 2002; Mazoyer et al., 2014). Also, this atypical organization is observed in some neurodevelopmental disorders such as schizophrenia, dyslexia, or autistic spectrum (Bishop, 2013; Sommer et al., 2001; Eyler et al., 2012).

How these atypical individuals organize the rest of their lateralized functions has been an extensively researched question. Historically, two different hypotheses have been proposed to explain the hemispheric specialization of the brain. The Causal hypothesis (Kosslyn, 1987; Hellige, 1990) argues that rapid activities requiring the implementation of sequences of cognitive processes are better performed from a single, unilateral control. As a result, these activities are innately programmed to be lateralized, and, according to this model, some functions will be performed better if they are controlled by different hemispheres. This evolutionary pressure engenders a complementary relationship between these lateralized functions. For example, language production predominantly lateralizes to the left hemisphere, while visuospatial attention typically favors the right hemisphere. As per the Causal hypothesis, if language were to atypically lateralize to the right hemisphere, visuospatial attention would, in turn, shift to the left hemisphere. The Statistical hypothesis (Bryden et al., 1983), on the other hand, postulates that each cognitive function lateralizes independently from the others. In this view, the likelihood of encountering atypical hemispheric dominance is essentially a statistical phenomenon, with no inherent interrelationship between various lateralized functions. The evidence supporting each theory in left-handers is mixed. Some studies have obtained data favoring the Statistical hypothesis when analyzing language, visuospatial attention, and face processing (Whitehouse and Bishop, 2009; Badzakova-Trajkov et al., 2010; Rosch et al., 2012). Most investigations, however, have demonstrated a mirrored organization of praxis (Vingerhoets et al., 2013; Gerrits et al., 2020), visuospatial attention (Badzakova-Trajkov et al., 2010; Gerrits et al., 2020; Cai et al., 2013), face recognition (Badzakova-Trajkov et al., 2010; Gerrits et al., 2020; Gerrits et al., 2019), and emotional prosody (Gerrits et al., 2020) in most individuals with atypical lateralization of language. Nonetheless, it should be noted that this reversed pattern is not observed in all atypical participants, and all the studies found specific cases in favor of both theories (Gerrits et al., 2020).

All these studies have employed a production task to investigate language lateralization in the inferior frontal cortex (IFC), classifying left-handed participants into typical and atypical groups. The rest of the lateralized cognitive functions have been examined using several tasks, but they all mainly depended on posterior parts of the brain. This may have limited the results, given that they were not dealing with directly homotopic functions, which would have been the most sensitive scenario to the proposed advantages of hemispheric specialization (Rogers, 2000; Vallortigara and Rogers, 2020). To directly address the language production function, which is typically dependent on the left IFC, we would have to use a cognitive task that is typically dependent on the homotopic right IFC. According to previous literature, this function would be inhibitory control. Extensive results support the notion that stopping an ongoing response involves a rightward network that includes the IFC, presupplementary motor area (preSMA), and subthalamic nucleus (STN) (Aron and Poldrack, 2006; Xue et al., 2008; Aron et al., 2014; Hannah and Aron, 2021). Thus, inhibitory control and language production together are the ideal candidates to investigate the Causal vs. Statistical disjunctive. But surprisingly, no investigation to date has analyzed this.

What causes language function to typically or atypically lateralize also remains unclear. Left-handedness appears to be, at best, a predisposing factor (Mazoyer et al., 2014). In clinical populations, atypical lateralization of language has been found to be a defining characteristic among patients of dyslexia (Bishop, 2013), schizophrenia (Sommer et al., 2001), and autism spectrum disorders (Eyler et al., 2012). However, aside from exhibiting poorer language performance (Mellet et al., 2014; Powell et al., 2012), no study to date has identified subclinical markers of these neurodevelopmental conditions among healthy individuals with atypical lateralization. Regarding the neural basis of language lateralization, the corpus callosum hypothesis (Gazzaniga, 2000)— which suggests that typical hemispheric specialization relies on the proper development of this structure — has garnered attention in recent years due to several noteworthy findings: (1) regional functional lateralization is directly linked to the size reduction of its interconnecting corpus callosum segments (Karolis et al., 2019); and (2) atypical lateralization of language has been associated with a larger corpus callosum and increased interhemispheric transfer at rest in language-relevant regions (Labache et al., 2020).

The primary aim of this study was to investigate the relationship between the lateralization of language production and inhibitory control. To achieve this, we divided non-right-handed participants into typically lateralized and atypically lateralized for language using the verb generation task. These two groups then completed the stop-signal task. In alignment with the predictions of the Causal hypothesis, we expected that: (1) the atypical group would exhibit a mirrored brain organization — particularly in the IFC, preSMA, and STN — during the stop-signal task; and (2) no individual would demonstrate lateralization of both language production and inhibitory control in the IFC of the same hemisphere. The secondary aim of this study was to explore certain potentially causal correlates of the lateralization of both functions. To test the presence of correlates supporting the corpus callosum hypothesis of lateralization, we analyzed the callosal volume and the interhemispheric functional connectivity at rest of the IFC, preSMA, and STN. To test the relation between atypical lateralization and several neurodevelopmental disorders (dyslexia, schizophrenia, and autism), we analyzed subclinical traits of these conditions among our healthy participants.

The present study employed the stop-signal task to investigate brain areas involved in the inhibitory control function in two different groups of individuals: typically and atypically lateralized for language production. As expected, the atypical participants showed, as a group, a mirrored brain organization of the inhibitory control function compared to the typical group. This leftward organization affected the entire inhibitory control network, including the IFC, preSMA, and STN. However, some participants manifested a clear overlap in the control of language and inhibitory functions. Our results also demonstrate that atypical organization of language production is associated with an increased white matter volume of the corpus callosum, and that atypical lateralization of inhibitory control is related with a higher interhemispheric functional coupling of the IFC. Behaviorally, atypical lateralization was not associated with performance on the task, but an association existed with worse reading accuracy, and more schizotypy and autistic traits.

Our first relevant result is that the group of individuals with atypical lateralization of language presented a mirrored brain organization during the stop-signal task compared to the typically lateralized group. Remarkably, the hemispheric asymmetry differences obtained affected the entire inhibitory control brain network (Hannah and Aron, 2021). The inferior frontal cortex, or IFC, associated with the initiation of the stop command and the main cortical area of this system, showed a clear lateralization effect, with the typical group almost exclusively using the right part and the atypical group mostly activating the left part. The presupplementary motor area, or pre-SMA, related to the implementation of the stop command, was more bilaterally involved in both groups, although atypicals activated the left part more and typicals activated the right part more. Importantly, the hemispheric reversal of organization also affected the subcortical structures participating in this network, namely the STN and thalamus. These structures displayed a lateralization pattern similar to the preSMA, with a more pronounced bilateral engagement but higher involvement of the right part in typicals and of the left part in atypicals. Thus, this is the first demonstration that an entire cerebral network controlling a cognitive function shows a completely flipped organization in an atypical population for language, which supports the Causal hypothesis of lateralization (Kosslyn, 1987; Hellige, 1990). Moreover, this result reveals that hemispheric lateralization goes beyond cerebral cortices, and even interhemispheric connectivity, through the corpus callosum, given that it also involves frontal-subcortical circuits. Additionally, the specificity of this result to the areas of the proposed network is another element that supports the relevance of this system in inhibitory control.

It should be noted that previous studies have demonstrated a significantly higher involvement of the left IFC during cognitive control tasks in bilingual individuals when compared to monolinguals (Garbin et al., 2010; Rodríguez-Pujadas et al., 2013). However, this could not be directly considered in the current study due to the limited number of monolinguals participants (n=8), which would not allow for statistically robust tests. So, we cannot rule out the possibility that bilingualism could be acting as an enabling mechanism among atypically lateralized participants for the shift of the inhibitory control network to the left hemisphere. Therefore, the findings of this study should be extrapolated to the monolingual population with caution.

Remarkably, this pattern of reversed specialization for inhibitory control was observed in the absence of a relation between functional lateralization and performance differences during the ‘go’ or ‘stop’ conditions. Even though the ‘go’ condition involves a lexical decision requiring language processing, and the ‘stop’ condition consists of inhibiting a linguistic process, the functional organization of the IFC during the tasks did not affect the response speed (RT and SSRT) or the ‘go’ accuracy in either case. Thus, brain organization for language and inhibition, as initially defined in our study, and cognitive efficiency are not directly related, based on our data. This finding is consistent with prior studies highlighting that: (1) a densely segregated functional organization, rather than a reversed organization, is linked to general (Gerrits et al., 2020) and specific (Mellet et al., 2014) cognitive deficits; and (2) specific cognitive deficits appear to be confined to the domains of verbal and spatial skills (Mellet et al., 2014).

Importantly, despite observing a complimentary relationship between language production and inhibitory control at the group level, not all the individuals presented a strong and segregated distribution of these two functions. The vast majority of hemispherically segregated individuals showed the typical segregation of language in the left hemisphere and inhibitory control in the right hemisphere, whereas only a low percentage displayed the opposite segregated organization. The rest, in agreement with similar reports on other functions (Gerrits et al., 2019), lacked a clear hemispheric lateralization of language production, inhibitory control, or both. In all cases, this implied some level of overlapping or sharing of the same area of the inferior frontal cortex. Both the low ratio of atypical segregation and the high ratio of bilateral inhibitory control could be explained by the fact that typical right hemispheric functions (such as inhibition) seem to be more bilaterally represented than typical left functions (such as language; Gotts et al., 2013). Some conclusions can be extracted from this: (1) Although infrequent, a strong atypical segregation of the two functions is possible; (2) Our data have not revealed any case with a segregation of both frontal cognitive functions in the same hemisphere; and (3) A large percentage of participants with a strong left lateralization of language presented a bilateral control of inhibition, showing that a typical dominance for language does not imply a strongly lateralized organization of other cognitive functions. The existence of exceptions to the group pattern of mirrored brain organization requires us to use caution when interpreting these data as supporting the causal hypothesis. Although the negative correlation between the lateralization indexes during the verb generation task and the stop-signal task is strong at the group level, some individual data support the statistical hypothesis (Bryden et al., 1983) instead. Given that we found no cases of both cognitive functions completely sharing a hemisphere (see Figure 2), we propose that causal-supporting vs. statistical-supporting results reflects two different pathways to cognitive control. On the one hand, the segregated pathway (in line with the Causal hypothesis) develops due to the cerebral bias towards lateralizing certain cognitive functions in a single hemisphere. On the other hand, the integrated pathway (in line with the Statistical hypothesis), appears when ontogenetic development towards strong lateralized control fails in some way.

These findings could have significant implications when studying neonatal lesions that facilitate the development of left-handedness (Dinomais et al., 2017), or when assessing the consequences of brain lesions in the left-handed population (Newport et al., 2022). For example, a left-sided lesion in an individual with rightward language lateralization may result in inhibitory control and social deficits (Mosch et al., 2005). It also raises questions about neuroplasticity, particularly how functional reorganization following a brain lesion (such as a left-to-right reorganization of language) would impact this hemispheric relationship between language and inhibitory control, and the cognitive consequences for inhibitory control after such a reorganization (Bates et al., 2001). These results could also offer valuable insights for cognitive rehabilitation procedures in patients with psychiatric disorders that seem to be linked to a higher prevalence of left-handedness and alterations in language lateralization (Webb et al., 2013).

The corpus callosum is the main cerebral structure in interhemispheric connectivity, with the genu area being responsible for the connectivity between the two inferior frontal gyri (Hofer and Frahm, 2006). Phylogenetic data have demonstrated a negative association between the corpus callosum volume and hemispheric specialization, suggesting that the ontogenetic development of the brain is supported by a decrease in interhemispheric connectivity to potentiate and establish intrahemispheric connectivity and hemispheric specialization (Tzourio-Mazoyer, 2016). There is evidence of this mechanism in language development (Tzourio-Mazoyer and Seghier, 2016) and in agenesis of the corpus callosum, which has been associated with lower lateralization of language and worse language performance (Hinkley et al., 2016; Bartha-Doering et al., 2021). In line with this proposal, we have replicated the relation between callosal enlargement and atypical functional organization found in a previous study (Labache et al., 2020). Moreover, in our data, this structural correlation was accompanied by an association between atypical lateralization of inhibitory control and strength of interhemispheric functional connectivity in the pars triangularis of the IFC.

As hypothesized, when considering the functional lateralization during both tasks, atypical organizations were found linearly related with higher scores on a schizotypy test (only for inhibitory control), higher scores on an autistic spectrum test (only for language production), and a higher rate of reading errors (both language production and inhibitory control). The first result is consistent with previous structural, functional, and behavioral data that have shown reduced language lateralization in schizophrenia patients (Sommer et al., 2001; Li et al., 2007). In the case of schizotypy, the evidence for reduced lateralization of cognitive functions is mixed and may respond to differences in the methodology employed (Park and Waldie, 2017). However, consistently with the present results, this condition has been previously associated to impaired behavioral and neural processing during the stop-signal task (Jia et al., 2021), and to an abnormal frontal functional asymmetry (Le et al., 2020). The second result relates atypical language lateralization with the presence of autistic traits, a result consistent with data obtained in autism (Jouravlev et al., 2020). The third result shows that atypical organization was related with a higher rate of reading errors when considering word length and word familiarity. The length effect has been considered a pathognomonic symptom of reading disorders such as developmental dyslexia (Provazza et al., 2019) and pure alexia (Roberts et al., 2013). These disorders have also been characterized by a weak language lateralization (Bishop, 2013). Thus, presented results support the hypothesis that atypical hemispheric specialization is related with worse cognitive performance in the linguistic domain, and even preclinical traits of some neurodevelopmental disorders among healthy population.

In conclusion, our study demonstrates a strong connection between the lateralization of language production in the IFC and the hemispheric specialization of the inhibitory control network. This functional interrelationship is apparent through both regional and voxel-wise analyses and impacts all the primary cerebral hubs of inhibitory control. Furthermore, the atypical lateralization of these two cognitive functions shows a linear correlation with increased inter-hemispheric functional connectivity in the IFC, a larger corpus callosum, the presence of subclinical markers of schizotypy and autism, and decreased reading accuracy. Consequently, our research provides compelling evidence in support of the causal and corpus callosum hypotheses of lateralization among left-handers, and offers valuable insights into the pathogenic mechanisms underlying atypical lateralization in healthy population.

All data presented in this study are included in figshare.

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   Structural (VBM), functional (fMRI) and behavioral data relative to the research article "What happens to the inhibitory control functions of the right inferior frontal cortex when this area is dominant for language?".

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Author details

1. Esteban Villar-Rodríguez

   Neuropsychology and Functional Neuroimaging; Jaume I University, Castellón de la Plana, Spain

   Contribution

   Data curation, Formal analysis, Investigation, Methodology, Writing – original draft, Writing – review and editing

   For correspondence

   esvillar@uji.es

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-9691-3776

2. Cristina Cano-Melle

   Neuropsychology and Functional Neuroimaging; Jaume I University, Castellón de la Plana, Spain

   Contribution

   Data curation, Investigation

   Competing interests

   No competing interests declared

3. Lidón Marin-Marin

   Neuropsychology and Functional Neuroimaging; Jaume I University, Castellón de la Plana, Spain

   Contribution

   Investigation

   Competing interests

   No competing interests declared

4. Maria Antònia Parcet

   Neuropsychology and Functional Neuroimaging; Jaume I University, Castellón de la Plana, Spain

   Contribution

   Conceptualization, Writing – review and editing

   Competing interests

   No competing interests declared

5. César Avila

   Neuropsychology and Functional Neuroimaging; Jaume I University, Castellón de la Plana, Spain

   Contribution

   Conceptualization, Supervision, Methodology, Writing – original draft, Project administration, Writing – review and editing

   For correspondence

   avila@psb.uji.es

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-5840-605X

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