To understand language, we must infer structured meanings from real-time auditory or visual signals. Researchers have long focused on word-by-word structure building in working memory as a mechanism that might enable this feat. However, some have argued that language processing does not typically involve rich word-by-word structure building, and/or that apparent working memory effects are underlyingly driven by surprisal (how predictable a word is in context). Consistent with this alternative, some recent behavioral studies of naturalistic language processing that control for surprisal have not shown clear working memory effects. In this fMRI study, we investigate a range of theory-driven predictors of word-by-word working memory demand during naturalistic language comprehension in humans of both sexes under rigorous surprisal controls. In addition, we address a related debate about whether the working memory mechanisms involved in language comprehension are language specialized or domain general. To do so, in each participant, we functionally localize (1) the language-selective network and (2) the "multiple-demand" network, which supports working memory across domains. Results show robust surprisal-independent effects of memory demand in the language network and no effect of memory demand in the multiple-demand network. Our findings thus support the view that language comprehension involves computationally demanding word-by-word structure building operations in working memory, in addition to any prediction-related mechanisms. Further, these memory operations appear to be primarily conducted by the same neural resources that store linguistic knowledge, with no evidence of involvement of brain regions known to support working memory across domains.SIGNIFICANCE STATEMENT This study uses fMRI to investigate signatures of working memory (WM) demand during naturalistic story listening, using a broad range of theoretically motivated estimates of WM demand. Results support a strong effect of WM demand in the brain that is distinct from effects of word predictability. Further, these WM demands register primarily in language-selective regions, rather than in "multiple-demand" regions that have previously been associated with WM in nonlinguistic domains. Our findings support a core role for WM in incremental language processing, using WM resources that are specialized for language.

Much research in cognitive neuroscience supports prediction as a canonical computation of cognition across domains. Is such predictive coding implemented by feedback from higher-order domain-general circuits, or is it locally implemented in domain-specific circuits? What information sources are used to generate these predictions? This study addresses these two questions in the context of language processing. We present fMRI evidence from a naturalistic comprehension paradigm (1) that predictive coding in the brain's response to language is domain-specific, and (2) that these predictions are sensitive both to local word co-occurrence patterns and to hierarchical structure. Using a recently developed continuous-time deconvolutional regression technique that supports data-driven hemodynamic response function discovery from continuous BOLD signal fluctuations in response to naturalistic stimuli, we found effects of prediction measures in the language network but not in the domain-general multiple-demand network, which supports executive control processes and has been previously implicated in language comprehension. Moreover, within the language network, surface-level and structural prediction effects were separable. The predictability effects in the language network were substantial, with the model capturing over 37% of explainable variance on held-out data. These findings indicate that human sentence processing mechanisms generate predictions about upcoming words using cognitive processes that are sensitive to hierarchical structure and specialized for language processing, rather than via feedback from high-level executive control mechanisms.

What role do domain-general executive functions play in human language comprehension? To address this question, we examine the relationship between behavioral measures of comprehension and neural activity in the domain-general "multiple demand" (MD) network, which has been linked to constructs like attention, working memory, inhibitory control, and selection, and implicated in diverse goal-directed behaviors. Specifically, functional magnetic resonance imaging data collected during naturalistic story listening are compared with theory-neutral measures of online comprehension difficulty and incremental processing load (reading times and eye-fixation durations). Critically, to ensure that variance in these measures is driven by features of the linguistic stimulus rather than reflecting participant- or trial-level variability, the neuroimaging and behavioral datasets were collected in nonoverlapping samples. We find no behavioral-neural link in functionally localized MD regions; instead, this link is found in the domain-specific, fronto-temporal "core language network," in both left-hemispheric areas and their right hemispheric homotopic areas. These results argue against strong involvement of domain-general executive circuits in language comprehension.