Linguistic Structure and Meaning Organize Neural Oscillations into a Content-Specific Hierarchy.

Neural oscillations track linguistic information during speech comprehension (Ding et al., 2016; Keitel et al., 2018), and are known to be modulated by acoustic landmarks and speech intelligibility (Doelling et al., 2014; Zoefel and VanRullen, 2015). However, studies investigating linguistic tracking have either relied on non-naturalistic isochronous stimuli or failed to fully control for prosody. Therefore, it is still unclear whether low-frequency activity tracks linguistic structure during natural speech, where linguistic structure does not follow such a palpable temporal pattern. Here, we measured electroencephalography (EEG) and manipulated the presence of semantic and syntactic information apart from the timescale of their occurrence, while carefully controlling for the acoustic-prosodic and lexical-semantic information in the signal. EEG was recorded while 29 adult native speakers (22 women, 7 men) listened to naturally spoken Dutch sentences, jabberwocky controls with morphemes and sentential prosody, word lists with lexical content but no phrase structure, and backward acoustically matched controls. Mutual information (MI) analysis revealed sensitivity to linguistic content: MI was highest for sentences at the phrasal (0.8-1.1 Hz) and lexical (1.9-2.8 Hz) timescales, suggesting that the delta-band is modulated by lexically driven combinatorial processing beyond prosody, and that linguistic content (i.e., structure and meaning) organizes neural oscillations beyond the timescale and rhythmicity of the stimulus. This pattern is consistent with neurophysiologically inspired models of language comprehension (Martin, 2016, 2020; Martin and Doumas, 2017) where oscillations encode endogenously generated linguistic content over and above exogenous or stimulus-driven timing and rhythm information.SIGNIFICANCE STATEMENT Biological systems like the brain encode their environment not only by reacting in a series of stimulus-driven responses, but by combining stimulus-driven information with endogenous, internally generated, inferential knowledge and meaning. Understanding language from speech is the human benchmark for this. Much research focuses on the purely stimulus-driven response, but here, we focus on the goal of language behavior: conveying structure and meaning. To that end, we use naturalistic stimuli that contrast acoustic-prosodic and lexical-semantic information to show that, during spoken language comprehension, oscillatory modulations reflect computations related to inferring structure and meaning from the acoustic signal. Our experiment provides the first evidence to date that compositional structure and meaning organize the oscillatory response, above and beyond prosodic and lexical controls.

Object shape and orientation do not routinely influence performance during language processing.

The role of visual representations during language processing remains unclear: They could be activated as a necessary part of the comprehension process, or they could be less crucial and influence performance in a task-dependent manner. In the present experiments, participants read sentences about an object. The sentences implied that the object had a specific shape or orientation. They then either named a picture of that object (Experiments 1 and 3) or decided whether the object had been mentioned in the sentence (Experiment 2). Orientation information did not reliably influence performance in any of the experiments. Shape representations influenced performance most strongly when participants were asked to compare a sentence with a picture or when they were explicitly asked to use mental imagery while reading the sentences. Thus, in contrast to previous claims, implied visual information often does not contribute substantially to the comprehension process during normal reading.

The contents of predictions in sentence comprehension: activation of the shape of objects before they are referred to.

When comprehending concrete words, listeners and readers can activate specific visual information such as the shape of the words' referents. In two experiments we examined whether such information can be activated in an anticipatory fashion. In Experiment 1, listeners' eye movements were tracked while they were listening to sentences that were predictive of a specific critical word (e.g., "moon" in "In 1969 Neil Armstrong was the first man to set foot on the moon"). 500 ms before the acoustic onset of the critical word, participants were shown four-object displays featuring three unrelated distractor objects and a critical object, which was either the target object (e.g., moon), an object with a similar shape (e.g., tomato), or an unrelated control object (e.g., rice). In a time window before shape information from the spoken target word could be retrieved, participants already tended to fixate both the target and the shape competitors more often than they fixated the control objects, indicating that they had anticipatorily activated the shape of the upcoming word's referent. This was confirmed in Experiment 2, which was an ERP experiment without picture displays. Participants listened to the same lead-in sentences as in Experiment 1. The sentence-final words corresponded to the predictable target, the shape competitor, or the unrelated control object (yielding, for instance, "In 1969 Neil Armstrong was the first man to set foot on the moon/tomato/rice"). N400 amplitude in response to the final words was significantly attenuated in the shape-related compared to the unrelated condition. Taken together, these results suggest that listeners can activate perceptual attributes of objects before they are referred to in an utterance.