Abstract
Language behaviour is complex, but neuroscientific evidence disentangles it into distinct components supported by dedicated brain areas or networks. In this Review, we describe the ‘core’ language network, which includes left-hemisphere frontal and temporal areas, and show that it is strongly interconnected, independent of input and output modalities, causally important for language and language-selective. We discuss evidence that this language network plausibly stores language knowledge and supports core linguistic computations related to accessing words and constructions from memory and combining them to interpret (decode) or generate (encode) linguistic messages. We emphasize that the language network works closely with, but is distinct from, both lower-level — perceptual and motor — mechanisms and higher-level systems of knowledge and reasoning. The perceptual and motor mechanisms process linguistic signals, but, in contrast to the language network, are sensitive only to these signals’ surface properties, not their meanings; the systems of knowledge and reasoning (such as the system that supports social reasoning) are sometimes engaged during language use but are not language-selective. This Review lays a foundation both for in-depth investigations of these different components of the language processing pipeline and for probing inter-component interactions.
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Data availability
The data used to generate the activation maps in Fig. 2a,b were released as part of Lipkin et al.49 and are available for download at https://figshare.com/articles/dataset/LanADataset/20425209. The data used to generate Fig. 2c were released as a supplement to Mahowald and Fedorenko51 and are available at: https://doi.org/10.6084/m9.figshare.22183564. The data used to generate the bar graphs in Figs. 2d and 3a,b come from published and pre-printed papers but, for convenience and ease of figure reproducibility, have been consolidated and placed on a dedicated Open Science Framework (OSF) page: https://osf.io/4tdcx/ (see Supplementary methods for details).
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Acknowledgements
The authors thank T. Gibson, W. Freiwald and N. Kanwisher for comments on the earlier drafts of the manuscript; A. Fung and C. Casto for help with figures and the Open Science Framework (OSF) page; A. Flinker, M. Long and J. Li for consultations on the anatomical locations of some functional areas; S. Swords, N. Jhingan, H. S. Kim and G. Tuckute for help with references; J. Gallée for help with creating the stimuli for the Broca’s area localizer (Fig. 4c); and S. Nastase for insightful and constructive comments. E.F. was supported by National Institutes of Health (NIH) awards R01-DC016607, R01-DC016950 and U01-NS121471, and by research funds from the McGovern Institute for Brain Research, the Brain and Cognitive Sciences Department, the Simons Center for the Social Brain and MIT’s Quest for Intelligence. A.A.I. was supported by funds from MIT’s Quest for Intelligence. T.I.R. was supported by a fellowship from the Poitras Center for Psychiatric Disorders Research at the McGovern Institute.
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Glossary
- Aphasia
-
Impairments in understanding and/or producing language as a result of brain damage (such as stroke or neural degeneration).
- Apraxia of speech
-
Impairments in producing sounds, syllables and words because of neurological problems with speech motor planning; speech of individuals with apraxia contains sound distortions, groping for sounds and errors in stress or rhythm, but they do not have difficulties making non-speech oral-motor movements.
- Causally important
-
For a particular function, if interfering with a neural unit’s (such as a cell or a brain area) activity or structural integrity leads to observable impairment of that function.
- Compositional semantics
-
The system of relationships between phrases and sentences and meanings; a key principle of compositional semantics in natural language is that the meaning of a multi-word sequence (for example, a phrase) is determined by the meanings of the composite words and the syntactic rules that were used to combine those words.
- Constructions
-
Learned pairings between a linguistic form and meaning; importantly, constructions encompass not only single morphemes and words but also multi-word sequences, which can correspond to complex meanings.
- Contrast
-
A pair of conditions that differ in a critical (stimulus-related or task-related) feature of interest; localizers use a particular contrast to localize a brain area or areas that support some perceptual, motor or cognitive function.
- Discourse
-
The system of relationships among clauses and sentences in multi-sentence sequences, including narratives and conversations.
- Double dissociations
-
Complementary cases of selective deficits in two perceptual, motor or cognitive functions (for example, if in one individual, language is impaired but general reasoning is preserved, but in another individual language is preserved but reasoning is impaired); or selective relationships between two brain areas and two perceptual, motor or cognitive functions where brain area A supports function F1 but not function F2, and brain area B supports function F2 but not F1 (for example, damage to the language areas leads to difficulties in understanding and producing language but leaves abilities supported by the multiple demand network, such as executive abilities and formal reasoning abilities, unimpaired; by contrast, damage to the multiple demand network areas leads to difficulties with executive control and reasoning but leaves linguistic functions unimpaired).
- Dysarthria of speech
-
Impairments in producing sounds, syllables and words because of weakening or improper coordination of the muscles of the articulatory organs (caused by brain or nerve damage); speech of individuals with dysarthria sounds slurred or mumbling and they also have difficulties in making non-speech oral-motor movements.
- Functional connectivity
-
Correlation in activity patterns between different brain areas, often measured during naturalistic cognition paradigms.
- Functional localization
-
An analytic approach that aggregates brain data from multiple participants while taking into account inter-individual differences in the precise locations of functional areas. This approach uses a localizer to find areas of interest within individual participants’ brains and then measures the response in these areas to some condition(s) of interest; group-level statistical comparisons are performed on the measures extracted from the individually identified areas, which circumvents the need to average brains.
- Group-averaging
-
An analytic approach that aggregates brain data from multiple participants by averaging individual brain images projected into a common coordinate space; because functional areas vary in their precise locations across individuals, this approach leads to blurring and can generate misleading results.
- Homotopic areas
-
Each brain area exists in two copies — one in each hemisphere; an area in one hemisphere that is the corresponding area in the other hemisphere (for example, the areas homotopic to the language areas are areas in the right hemisphere that correspond to the left-hemisphere language areas).
- Language processing pipeline
-
A set of perceptual, motor and cognitive processes that jointly enable language comprehension or language production: in comprehension, the pipeline encompasses perceptual processing of linguistic inputs, mapping linguistic forms to meanings and integrating these meanings with preceding linguistic context and non-linguistic knowledge sources; in production, the pipeline encompasses the transformation of intended meanings into linguistic form, and planning and generating the physical output.
- Lexical semantics
-
The system of relationships between word forms and word meanings.
- Linguistic regularities
-
Structure, or regular patterns, in linguistic sequences; regular patterns characterize all levels of language, from sounds to words, to sentences, to connected discourses.
- Meanings
-
The meanings of words and phrases (for example, the word ‘dog’ has meaning to an English speaker, whereas the word ‘chien’ (‘dog’ in French) or a non-word ‘xog’ does not have meaning to an English speaker, under typical circumstances); or a collection of associative, abstract and generalizable knowledge associated with a given cue, either linguistic or non-linguistic (for example, the word ‘dog’, a picture of a dog and the sound of a dog’s barking are all associated with a similar meaning, although different cues or contexts may make some aspects of the meaning more or less salient).
- Modality-independent
-
If a brain area responds to stimuli across different input or output modalities (for example, spoken language, written language or signed language).
- Morphology
-
The system of relationships among ‘morphemes’, which are the smallest meaning-bearing units in a language; morphemes can be complete words, such as ‘cat’, but also word parts, such as ‘-ful’ and ‘anti-’.
- Naturalistic cognition paradigms
-
Paradigms that do not rely on controlled, experimenter-crafted conditions and contrasts but, instead, present rich naturalistic stimuli (such as movies or narratives) or feature open-ended designs, such as a resting state (task-free, stimulus-free periods during the study).
- Phonology
-
The system of relationships among speech sounds in a language; the rules that govern the possible combinations and orderings of sounds are called phonotactics.
- Single dissociation
-
A selective deficit in a perceptual, motor or cognitive function (for example, aphasia is a selective deficit in understanding or producing language); or a selective relationship between some brain area and a perceptual, motor or cognitive function (for example, brain damage to the language areas leads to aphasia but leaves other cognitive processes unimpaired).
- Surface properties of linguistic stimuli
-
Properties that are tied to the form of a linguistic stimulus; the form is determined by the input or output modality (for example, for speech they have to do with the speech sounds, and for written language they have to do with the visual shapes of letters).
- Syntax
-
The system of constraints on how words can combine into phrases and sentences to create complex meanings.
- Tasks
-
What participants are asked to do during an experiment (for example, passive reading or answering comprehension questions).
- Theory of mind
-
The ability to understand and take into account another individual’s mental state.
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Fedorenko, E., Ivanova, A.A. & Regev, T.I. The language network as a natural kind within the broader landscape of the human brain. Nat. Rev. Neurosci. 25, 289–312 (2024). https://doi.org/10.1038/s41583-024-00802-4
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DOI: https://doi.org/10.1038/s41583-024-00802-4
