Angela D. Friederici

Learning takes place throughout lifetime but differs in infants and adults because of the development of the PFC, a brain region responsible for cognitive control. To test this hypothesis, adults were investigated in a language learning paradigm under inhibitory, cathodal transcranial direct current stimulation over PFC. The experiment included a learning session interspersed with test phases and a test-only session. The stimulus material required the learning of grammatical dependencies between two elements in a novel language. In a parallel design, cathodal transcranial direct current stimulation over the left PFC, right PFC, or sham stimulation was applied during the learning session but not during the test-only session. Event-related brain potentials (ERPs) were recorded during both sessions. Whereas no ERP learning effects were observed during the learning session, different ERP learning effects as a function of prior stimulation type were found during the test-only session, although behavioral learning success was equal across conditions. With sham stimulation, the ERP learning effect was reflected in a centro-parietal N400-like negativity indicating lexical processes. Inhibitory stimulation over the left PFC, but not over the right PFC, led to a late positivity similar to that previously observed in prelinguistic infants indicating associative learning. The present data demonstrate that adults can learn with and without cognitive control using different learning mechanisms. In the presence of cognitive control, adult language learning is lexically guided, whereas it appears to be associative in nature when PFC control is downregulated.

We aimed to dissociate two levels of hierarchical structure building in language and mathematics, namely ‘first-level’ (the build-up of hierarchical structure with externally given elements) and ‘second-level’ (the build-up of hierarchical structure with internally represented elements produced by first-level processes). Using functional magnetic resonance imaging, we investigated these processes in three domains: sentence comprehension, arithmetic calculation (using Reverse Polish notation, which gives two operands followed by an operator) and a working memory control task. All tasks required the build-up of hierarchical structures at the first- and second-level, resulting in a similar computational hierarchy across language and mathematics, as well as in a working memory control task. Using a novel method that estimates the difference in the integration cost for conditions of different trial durations, we found an anterior-to-posterior functional organization in the prefrontal cortex, according to the level of hierarchy. Common to all domains, the ventral premotor cortex (PMv) supports first-level hierarchy building, while the dorsal pars opercularis (POd) subserves second-level hierarchy building, with lower activation for language compared with the other two tasks. These results suggest that the POd and the PMv support domain-general mechanisms for hierarchical structure building, with the POd being uniquely efficient for language.

Language processing is a trait of human species. The knowledge about its neurobiological basis has been increased considerably over the past decades. Different brain regions in the left and right hemisphere have been identified to support particular language functions. Networks involving the temporal cortex and the inferior frontal cortex with a clear left lateralization were shown to support syntactic processes, whereas less lateralized temporo-frontal networks subserve semantic processes. These networks have been substantiated both by functional as well as by structural connectivity data. Electrophysiological measures indicate that within these networks syntactic processes of local structure building precede the assignment of grammatical and semantic relations in a sentence. Suprasegmental prosodic information overtly available in the acoustic language input is processed predominantly in a temporo-frontal network in the right hemisphere associated with a clear electrophysiological marker. Studies with patients suffering from lesions in the corpus callosum reveal that the posterior portion of this structure plays a crucial role in the interaction of syntactic and prosodic information during language processing.

The ability to learn language is a human trait. In adults and children, brain imaging studies have shown that auditory language activates a bilateral frontotemporal network with a left hemispheric dominance. It is an open question whether these activations represent the complete neural basis for language present at birth. Here we demonstrate that in 2-d-old infants, the language-related neural substrate is fully active in both hemispheres with a preponderance in the right auditory cortex. Functional and structural connectivities within this neural network, however, are immature, with strong connectivities only between the two hemispheres, contrasting with the adult pattern of prevalent intrahemispheric connectivities. Thus, although the brain responds to spoken language already at birth, thereby providing a strong biological basis to acquire language, progressive maturation of intrahemispheric functional connectivity is yet to be established with language exposure as the brain develops.

Although the neural network for language processing in the adult brain is well specified, the neural underpinning of language acquisition is still underdetermined. Here, we define the milestones of syntax acquisition and discuss the possible neurophysiological preconditions thereof. Early language learning seems to be based on the bilateral temporal cortices. Subsequent syntax acquisition apparently primarily recruits a neural network involving the left frontal cortex and the temporal cortex connected by a ventrally located fiber system. The late developing ability to comprehend syntactically complex sentences appears to require a neural network that connects Broca’s area to the left posterior temporal cortex via a dorsally located fiber pathway. Thus, acquisition of syntax requires the maturation of fiber bundles connecting the classical language-relevant brain regions.