Review
The biological basis of language: insight from developmental grammatical impairments

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Highlights

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    Specific language impairment is a heterogeneous family of genetic developmental disorders.

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    We have identified a subtype, Grammatical-SLI, which affects the children's syntax, morphology, and phonology in similar ways.

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    Grammatical abilities are not impaired across the board: the children handle forms that are local, linear, semantic, and holistic, while stumbling on those that are nonlocal, hierarchical, abstract, and composed.

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    The mosaic of impaired and spared abilities is consistent with new models of the neural bases of syntax, morphology, and phonology which distinguish several dorsal and ventral language pathways in the brain.

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    We foresee substantial progress in the biology of language – evolution, genetics, neurobiology, computation, behavior – if language and language impairments are differentiated into underlying pathways and components.

Specific language impairment (SLI), a genetic developmental disorder, offers insights into the neurobiological and computational organization of language. A subtype, Grammatical-SLI (G-SLI), involves greater impairments in ‘extended’ grammatical representations, which are nonlocal, hierarchical, abstract, and composed, than in ‘basic’ ones, which are local, linear, semantic, and holistic. This distinction is seen in syntax, morphology, and phonology, and may be tied to abnormalities in the left hemisphere and basal ganglia, consistent with new models of the neurobiology of language which distinguish dorsal and ventral processing streams. Delineating neurolinguistic phenotypes promises a better understanding of the effects of genes on the brain circuitry underlying normal and impaired language abilities.

Section snippets

Developmental disorders as a window into the biology of language

Given the lack of animal models for language, and the inability to use invasive procedures with humans except out of medical necessity, our knowledge of the neurobiology of language has long depended upon natural experiments. During the 19th and 20th centuries, studies of patients with acquired brain lesions provided key insights 1, 2, 3. Understanding of language in the 21st century promises to be enriched by data from developmental disorders. SLI, a family of language impairments in otherwise

Grammatical phenotypes of SLI

This review focuses on what G-SLI can reveal about the structure and neural instantiation of language. Crucially, G-SLI is not a global impairment of language or even of grammar, but is strongly manifested in certain aspects of linguistic performance while leaving others largely intact. This raises the possibility that the contrast reflects a key division within the neural or genetic substrates of language. In particular, children with G-SLI have difficulty interpreting and producing syntactic

Extended versus Basic syntax

Extended syntax involves hierarchical structures and dependencies between words, often spanning the entire clause, which are computed in real time. For example, in ‘wh’-questions such as ‘Who did Joe see __?’, the ‘wh’-word and the empty position after ‘see’ are in a dependent relation, which may be analyzed as the movement of the word from its original position in an underlying structure (Figure 1). Additionally, Extended syntax is abstract: the assembly and interpretation of phrases depend on

Extended versus Basic morphology

G-SLI also impairs morphology in a selective fashion which targets abstract combinatorial operations. Extended morphological objects are generated, perceived, and represented as combinations of two or more morphemes, whereas Basic ones are stored and retrieved as wholes, or at best as containing parts which do not freely combine with other parts 9, 10. For example, in English, irregular past-tense forms (‘ate’, ‘bought’) are basic and handled by memory, whereas regular past-tense forms

Neural substrates of the distinction between Basic and Extended grammar

Recent advances using electroencephalography (EEG), magnetoencephalography (MEG), functional MRI (fMRI), and diffusion tensor imaging (DTI) tractography have led to sophisticated new models of the neural organization of language 35, 43, 44, 45. They allow us to assess whether the Extended processes, which pose the greatest problems in G-SLI, implicate brain networks that are different from those supporting Basic processes.

Neuroanatomy

A recent meta-analysis of 25 studies of the neuroanatomical correlates of broad SLI revealed consistent structural anomalies in two regions: the frontal cortex, particularly Broca's area and its right-hemisphere homologue, and the caudate nucleus [56]. The findings are generally consistent with SLI being a deficit of Extended processing (though Broca's area participates in aspects of Basic syntax and probably Basic morphology as well). The heterogeneity of SLI may mean that any meta-analysis of

Concluding remarks

We have suggested that the study of language impairment has opened a new frontier of research which promises a comprehensive biological understanding of language, from evolution and genes, through neuroanatomy and neural function, to linguistic computation, and thence to overt speech and comprehension. These advances depend on a multi-level approach. Rather than mapping genetic variants directly to overall language impairment, researchers must characterize the intermediate links by probing for

Acknowledgments

This paper is a capstone to the research of Heather van der Lely, who died of cancer in February, 2014. She identified the grammatical subtype of SLI, characterized it empirically (with collaborators in London, Paris, Louvain la Neuve, and Berlin), proposed the Extended–Basic distinction to explain the pattern of deficits, and wrote the first draft of this paper. Steven Pinker rewrote and fact-checked the paper before and after Heather's death, with extensive input from her main collaborator

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