Fractionating difficulty during sentence comprehension using functional neuroimaging.

Sentence comprehension is highly practiced and largely automatic, but this belies the complexity of the underlying processes. We used functional neuroimaging to investigate garden-path sentences that cause difficulty during comprehension, in order to unpack the different processes used to support sentence interpretation. By investigating garden-path and other types of sentences within the same individuals, we functionally profiled different regions within the temporal and frontal cortices in the left hemisphere. The results revealed that different aspects of comprehension difficulty are handled by left posterior temporal, left anterior temporal, ventral left frontal, and dorsal left frontal cortices. The functional profiles of these regions likely lie along a spectrum of specificity to generality, including language-specific processing of linguistic representations, more general conflict resolution processes operating over linguistic representations, and processes for handling difficulty in general. These findings suggest that difficulty is not unitary and that there is a role for a variety of linguistic and non-linguistic processes in supporting comprehension.

Updating constructions: additive effects of prior and current experience during sentence production.

While much earlier work has indicated that prior verb bias from lifelong language experience influences language processing, recent findings highlight the fact that verb biases induced during lab-based exposure sessions also influence processing. We investigated the nature of updating, i.e., how prior and current experience might interact in guiding subsequent sentence production. Participants underwent a short training session where we manipulated the bias of known English dative verbs. The prior bias of each verb for the double-object (DO) versus the prepositional-object (PO) dative was estimated using a corpus. Current verb bias was counterbalanced and controlled experimentally. Bayesian mixed-effects logistic models of participants' responses (DO or PO) during subsequent free-choice production showed that both the prior and current verb biases affected speakers' construction choice. These effects were additive and not interactive, contrary to the prediction from error-based learning models. Semantic similarity to other verbs and their experimentally manipulated biases influenced sentence production, consistent with item-based analogy and exemplar theory. These results shed light on the potential mechanisms underlying language updating and the adaptation of sentence production to ongoing experience.

Multimodal measures of sentence comprehension in agrammatism.

Agrammatic or asyntactic comprehension is a common language impairment in aphasia. We considered three possible hypotheses about the underlying cause of this deficit, namely problems in syntactic processing, over-reliance on semantics, and a deficit in cognitive control. We tested four individuals showing asyntactic comprehension on their comprehension of syntax-semantics conflict sentences (e.g., The robber handcuffed the cop), where semantic cues pushed towards a different interpretation from syntax. Two of the four participants performed above chance on such sentences indicating that not all agrammatic individuals are impaired in structure-based interpretation. We collected additional eyetracking measures from the other two participants, who performed at chance on the conflict sentences. These measures suggested distinct underlying processing profiles in the two individuals. Cognitive assessments further suggested that one participant might have performed poorly due to a linguistic cognitive control impairment while the other had difficulty due to over-reliance on semantics. Together, the results highlight the importance of multimodal measures for teasing apart aphasic individuals' underlying deficits. They corroborate findings from neurotypical adults by showing that semantics can strongly influence comprehension and that cognitive control could be relevant for choosing between competing sentence interpretations. They extend previous findings by demonstrating variability between individuals with aphasia-cognitive control might be especially relevant for patients who are not overly reliant on semantics. Clinically, the identification of distinct underlying problems in different individuals suggests that different treatment paths might be warranted for cases who might look similar on behavioral assessments.

Variation in how cognitive control modulates sentence processing.

Prior research suggests that cognitive control can assist the comprehension of sentences that create conflict between interpretations, at least under some circumstances. However, the mixed pattern of results suggests that cognitive control may not always be necessary for accurate comprehension. We tested whether cognitive control recruitment for language processing is systematically variable, depending on the type of sentential ambiguity or conflict, individual differences in cognitive control, and task demands. Participants completed two sessions in a web-based experiment. The first session tested conflict modulation using interleaved Stroop and sentence comprehension trials. Critical sentences contained syntax-semantics or phrase-attachment conflict. In the second session, participants completed three cognitive control and three working memory tasks. Exploratory factor analysis was used to index individual differences in a cognitive control factor and a working memory factor. At the group level, there were no significant conflict modulation effects for either syntax-semantics or phrase-attachment conflict. At the individual differences level, the cognitive control factor correlated with offline comprehension accuracy but not online processing measures for both types of conflict. Together, the results suggest that the role of cognitive control in sentence processing may vary according to task demands. When overt decisions are required, individual differences in cognitive control may matter such that better cognitive control results in better language comprehension performance. The results add to the mixed evidence on conflict modulation and raise questions about the situations under which cognitive control influences online processing.

Rational and Flexible Adaptation of Sentence Production to Ongoing Language Experience.

Whether sentences are formulated primarily using lexically based or non-lexically based information has been much debated. In this perspective article, I review evidence for rational flexibility in the sentence production architecture. Sentences can be constructed flexibly via lexically dependent or independent routes, and rationally depending on the statistical properties of the input and the validity of lexical vs. abstract cues for predicting sentence structure. Different neural pathways appear to be recruited for individuals with different executive function abilities and for verbs with different statistical properties, suggesting that alternative routes are available for producing the same structure. Together, extant evidence indicates that the human brain adapts to ongoing language experience during adulthood, and that the nature of the adjustment may depend rationally on the statistical contingencies of the current context.

Distributional learning in English: The effect of verb-specific biases and verb-general semantic mappings on sentence production.

Previous studies using artificial languages suggest that sentence production can be guided by verb-specific as well as verb-general statistics present in the language input. Here we investigated whether the statistical properties of ongoing input in the speakers' native language systematically affected their sentence production. Three experiments used a training-plus production test paradigm to examine whether sentence production comes to rely more on verb-specific biases or on verb-general semantic mappings depending on the relative predictive validities of those cues in the input. We found different patterns in speakers' structural choices and utterance characteristics depending on the relative validity of different cues. These results show that native language production can be updated by new input and suggest that the nature of the updating can be explained at least in part by adopting a cue-validity approach. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Neural Mechanisms Underlying the Dynamic Updating of Native Language.

Language users encounter different sentence structures from different people in different contexts. Although syntactic variability and adults' ability to adapt to it are both widely acknowledged, the relevant mechanisms and neural substrates are unknown. We hypothesized that syntactic updating might rely on cognitive control, which can help detect and resolve mismatch between prior linguistic expectations and new language experiences that countervail those expectations and thereby assist in accurately encoding new input. Using functional neuroimaging (fMRI), we investigated updating in garden-path sentence comprehension to test the prediction that regions within the left inferior frontal cortex might be relevant neural substrates, and additionally, explored the role of regions within the multiple demand network. Participants read ambiguous and unambiguous main-verb and relative-clause sentences. Ambiguous relative-clause sentences led to a garden-path effect in the left pars opercularis within the lateral frontal cortex and the left anterior insula/frontal operculum within the multiple demand network. This effect decreased upon repeated exposure to relative-clause sentences, consistent with updating. The two regions showed several contrastive patterns, including different activation relative to baseline, correlation with performance in a cognitive control task (the Stroop task), and verb-specificity versus generality in adaptation. Together, these results offer new insight into how the brain updates native language. They demonstrate the involvement of left frontal brain regions in helping the language system adjust to new experiences, with different areas playing distinct functional roles.

Who did what? A causal role for cognitive control in thematic role assignment during sentence comprehension.

Thematic role assignment - generally, figuring out who did what to whom - is a critical component of sentence comprehension, which is influenced by both syntactic and semantic cues. Conflict between these cues can result in temporary consideration of multiple incompatible interpretations during real-time sentence processing. We tested whether the resolution of syntax-semantics conflict can be expedited by the online engagement of cognitive control processes that are routinely used to regulate behavior across domains. In this study, cognitive control deployment from a previous Stroop trial influenced eye movements during subsequent sentence comprehension. Specifically, when syntactic and semantic cues competed for influence on interpretation, dynamic cognitive control engagement led to (a) fewer overall looks to a picture illustrating the competing but incorrect interpretation (Experiment 1), or (b) steeper growth in looks to a picture illustrating the correct interpretation (Experiment 2). Thus, prior cognitive control engagement facilitated the resolution of syntax-semantics conflict by biasing processing towards the intended analysis. This conflict adaptation effect demonstrates a causal connection between cognitive control and real-time thematic role assignment. Broader patterns demonstrated that prior cognitive control engagement also modulated sentence processing irrespective of the presence of conflict, reflecting increased integration of newly arriving cues with prior sentential content. Together, the results suggest that cognitive control helps listeners determine correct event roles during real-time comprehension.

Statistical experience and individual cognitive differences modulate neural activity during sentence production.

I explored how individual cognitive differences combine with prior statistical experience to determine choice of sentence structure during speech. Participants were exposed to English language input with controlled statistical properties wherein some verbs appeared equally often in two possible structures and others appeared in only one. Subsequently, they produced sentences naturally while their brain activity was scanned. Choosing a less preferred over a more preferred structure recruited regions involved in conflict control, especially in individuals with better control abilities. Activity within a key region, the anterior cingulate cortex or ACC, varied parametrically with the statistical input properties. ACC activation showed different correlations with language production and different functional connectivity patterns for different verbs. These results demonstrate how the adult brain adjusts to ongoing language experience and recruits different neural resources to accomplish the same speech goal under different circumstances.

Verb bias and verb-specific competition effects on sentence production.

How do speakers choose between structural options for expressing a given meaning? Overall preference for some structures over others as well as prior statistical association between specific verbs and sentence structures ("verb bias") are known to broadly influence language use. However, the effects of prior statistical experience on the planning and execution of utterances and the mechanisms that facilitate structural choice for verbs with different biases have not been fully explored. In this study, we manipulated verb bias for English double-object (DO) and prepositional-object (PO) dative structures: some verbs appeared solely in the DO structure (DO-only), others solely in PO (PO-only) and yet others equally in both (Equi). Structural choices during subsequent free-choice sentence production revealed the expected dispreference for DO overall but critically also a reliable linear trend in DO production that was consistent with verb bias (DO-only > Equi > PO-only). Going beyond the general verb bias effect, three results suggested that Equi verbs, which were associated equally with the two structures, engendered verb-specific competition and required additional resources for choosing the dispreferred DO structure. First, DO production with Equi verbs but not the other verbs correlated with participants' inhibition ability. Second, utterance duration prior to the choice of a DO structure showed a quadratic trend (DO-only < Equi > PO-only) with the longest durations for Equi verbs. Third, eye movements consistent with reimagining the event also showed a quadratic trend (DO-only < Equi > PO-only) prior to choosing DO, suggesting that participants used such recall particularly for Equi verbs. Together, these analyses of structural choices, utterance durations, eye movements and individual differences in executive functions shed light on the effects of verb bias and verb-specific competition on sentence production and the role of different executive functions in choosing between sentence structures.

Cognitive control during sentence generation.

Sentence generation involves selecting the right words out of multiple co-activated representations for the right utterance positions. This selection process is hypothesized to involve cognitive control. Previous research has linked cognitive control during language processing to regions within the frontal lobe, including the anterior cingulate cortex (ACC) and left ventrolateral prefrontal cortex (VLPFC). We examined the neural correlates of word and sentence generation to determine whether these cognitive control regions were activated more for sentences than words. We also manipulated the phonological overlap between nouns in the sentence to determine whether cognitive control recruitment increased under phonological competition. Whole-brain analysis revealed increased activation in the ACC and bilateral frontal cortex for sentences over words. Region of interest analysis revealed an effect of phonological competition within the ACC. Psychophysiological interaction (PPI) analysis further showed that functional connectivity within a task-maintenance network increased under phonological competition. These results suggest a role for conflict detection and monitoring during sentence generation, which is consistent with a theoretical framework incorporating domain-general cognitive control processes into language processing.

Acquiring and Producing Sentences: Whether Learners Use Verb-Specific or Verb-General Information Depends on Cue Validity.

Learning to produce sentences involves learning patterns that enable the generation of new utterances. Language contains both verb-specific and verb-general regularities that are relevant to this capacity. Previous research has focused on whether one source is more important than the other. We tested whether the production system can flexibly learn to use either source, depending on the predictive validity of different cues in the input. Participants learned new sentence structures in a miniature language paradigm. In three experiments, we manipulated whether individual verbs or verb-general mappings better predicted the structures heard during learning. Evaluation of participants' subsequent production revealed that they could use either the structural preferences of individual verbs or abstract meaning-to-form mappings to construct new sentences. Further, this choice varied according to cue validity. These results demonstrate flexibility within the production architecture and the importance of considering how language was learned when discussing how language is used.

Ventral and dorsal streams for choosing word order during sentence production.

Proficient language use requires speakers to vary word order and choose between different ways of expressing the same meaning. Prior statistical associations between individual verbs and different word orders are known to influence speakers' choices, but the underlying neural mechanisms are unknown. Here we show that distinct neural pathways are used for verbs with different statistical associations. We manipulated statistical experience by training participants in a language containing novel verbs and two alternative word orders (agent-before-patient, AP; patient-before-agent, PA). Some verbs appeared exclusively in AP, others exclusively in PA, and yet others in both orders. Subsequently, we used sparse sampling neuroimaging to examine the neural substrates as participants generated new sentences in the scanner. Behaviorally, participants showed an overall preference for AP order, but also increased PA order for verbs experienced in that order, reflecting statistical learning. Functional activation and connectivity analyses revealed distinct networks underlying the increased PA production. Verbs experienced in both orders during training preferentially recruited a ventral stream, indicating the use of conceptual processing for mapping meaning to word order. In contrast, verbs experienced solely in PA order recruited dorsal pathways, indicating the use of selective attention and sensorimotor integration for choosing words in the right order. These results show that the brain tracks the structural associations of individual verbs and that the same structural output may be achieved via ventral or dorsal streams, depending on the type of regularities in the input.

Controlled processing during sequencing.

Longstanding evidence has identified a role for the frontal cortex in sequencing within both linguistic and non-linguistic domains. More recently, neuropsychological studies have suggested a specific role for the left premotor-prefrontal junction (BA 44/6) in selection between competing alternatives during sequencing. In this study, we used neuroimaging with healthy adults to confirm and extend knowledge about the neural correlates of sequencing. Participants reproduced visually presented sequences of syllables and words using manual button presses. Items in the sequence were presented either consecutively or concurrently. Concurrent presentation is known to trigger the planning of multiple responses, which might compete with one another. Therefore, we hypothesized that regions involved in controlled processing would show greater recruitment during the concurrent than the consecutive condition. Whole-brain analysis showed concurrent > consecutive activation in sensory, motor and somatosensory cortices and notably also in rostral-dorsal anterior cingulate cortex. Region of interest analyses showed increased activation within left BA 44/6 and correlation between this region's activation and behavioral response times. Functional connectivity analysis revealed increased connectivity between left BA 44/6 and the posterior lobe of the cerebellum during the concurrent than the consecutive condition. These results corroborate recent evidence and demonstrate the involvement of BA 44/6 and other control regions when ordering co-activated representations.

Subdivision of frontal cortex mechanisms for language production in aphasia.

Ventrolateral prefrontal cortex (VLPFC) has long been linked to language production, but the precise mechanisms are still being elucidated. Using neuropsychological case studies, we explored possible sub-specialization within this region for different linguistic and executive functions. Frontal patients with different lesion profiles completed two sequencing tasks, which were hypothesized to engage partially overlapping components. The multi-word priming task tested the sequencing of co-activated representations and the overriding of primed word orders. The sequence reproduction task tested the sequencing of co-activated representations, but did not employ a priming manipulation. We compared patients' performance on the two tasks to that of healthy, age-matched controls. Results are partially consistent with an anterior-posterior gradient of cognitive control within lateral prefrontal cortex (Koechlin & Summerfield, 2007). However, we also found a stimulus-specific pattern, which suggests that sub-specialization might be contingent on type of representation as well as type of control signal. Isolating such components functionally and anatomically might lead to a better understanding of language production deficits in aphasia.

Parametric effects of syntactic-semantic conflict in Broca's area during sentence processing.

The hypothesized role of Broca's area in sentence processing ranges from domain-general executive function to domain-specific computation that is specific to certain syntactic structures. We examined this issue by manipulating syntactic structure and conflict between syntactic and semantic cues in a sentence processing task. Functional neuroimaging revealed that activation within several Broca's area regions of interest reflected the parametric variation in syntactic-semantic conflict. These results suggest that Broca's area supports sentence processing by mediating between multiple incompatible constraints on sentence interpretation, consistent with this area's well-known role in conflict resolution in other linguistic and non-linguistic tasks.

The neural basis of reversible sentence comprehension: evidence from voxel-based lesion symptom mapping in aphasia.

We explored the neural basis of reversible sentence comprehension in a large group of aphasic patients (n = 79). Voxel-based lesion symptom mapping revealed a significant association between damage in temporo-parietal cortex and impaired sentence comprehension. This association remained after we controlled for phonological working memory. We hypothesize that this region plays an important role in the thematic or what-where processing of sentences. In contrast, we detected weak or no association between reversible sentence comprehension and the ventrolateral pFC, which includes Broca's area, even for syntactically complex sentences. This casts doubt on theories that presuppose a critical role for this region in syntactic computations.

Selection for position: the role of left ventrolateral prefrontal cortex in sequencing language.

Patients with damage involving left ventrolateral prefrontal cortex (left VLPFC) often show syntactic deficits. They also show exaggerated interference effects during a variety of non-syntactic tasks, including picture naming and working memory. Conceivably, both deficits could arise from inadequate biasing of competitive interactions during language production. To test this hypothesis, we manipulated "positional" interference during multi-word naming by priming one of the nouns in the same or different position. Experimental case studies of four left VLPFC patients revealed that two of the patients showed exaggerated positional interference, greater number of errors, including omissions during multi-word production, increased production difficulty when the order of nouns did not match the predominant English pattern, as well as impaired comprehension of non-canonical reversible sentences. These results suggest that these two patients had an impairment in "selection for position". Different from the other two, their lesions included a subregion of frontal cortex (BA 44/6) that has been shown in neuroimaging studies to play a role in sequencing.

Give and take: syntactic priming during spoken language comprehension.

Syntactic priming during language production is pervasive and well-studied. Hearing, reading, speaking or writing a sentence with a given structure increases the probability of subsequently producing the same structure, regardless of whether the prime and target share lexical content. In contrast, syntactic priming during comprehension has proven more elusive, fueling claims that comprehension is less dependent on general syntactic representations and more dependent on lexical knowledge. In three experiments we explored syntactic priming during spoken language comprehension. Participants acted out double-object (DO) or prepositional-object (PO) dative sentences while their eye movements were recorded. Prime sentences used different verbs and nouns than the target sentences. In target sentences, the onset of the direct-object noun was consistent with both an animate recipient and an inanimate theme, creating a temporary ambiguity in the argument structure of the verb (DO e.g., Show the horse the book; PO e.g., Show the horn to the dog). We measured the difference in looks to the potential recipient and the potential theme during the ambiguous interval. In all experiments, participants who heard DO primes showed a greater preference for the recipient over the theme than those who heard PO primes, demonstrating across-verb priming during online language comprehension. These results accord with priming found in production studies, indicating a role for abstract structural information during comprehension as well as production.