Mary has a little chair: Eliciting noun-modifier phrases in individuals with acute post-stroke aphasia.

Background: Aphasia assessment primarily examines an individual's syntax, nouns, and verbs. However, modifiers, such as adjectives and number words, and bound morphemes can be the subject of considerable difficulty for individuals with aphasia. The Morphosyntactic Generation (MorGen) targets nouns, modifiers, and bound inflectional morphemes in two-word phrases among people with aphasia. Aims: The purpose of this work is to provide the first report of the MorGen in hyperacute-acute aphasia. In doing so, we aim to (1) examine the MorGen's concurrent validity with common assessments of aphasia; (2) describe performance in modifiers by people with acute aphasia; and (3) associate MorGen performance with extent of lesioned vascular territories in acute stroke. Methods & Procedures: 62 adult English speakers within the first 14 days of left hemisphere ischemic stroke and 61 healthy control participants completed the MorGen. In addition to receiving the MorGen, participants with stroke received the Western Aphasia Battery (WAB), Boston Naming Test, and Hopkins Action Naming Assessment. Clinical MRIs were analyzed for the extent of lesion in the vascular territory of the left anterior, medial, and posterior cerebral artery, as well as the left posterior choroidal and thalamoperforator arteries. Outcomes & Results: Aim 1: Performance on the MorGen demonstrated consistently high, significant correlations with that on the WAB, Boston Naming Test, and Hopkins Action Naming Assessment. Aim 2: Individuals who had a stroke but were within functional limits (WFL) on the WAB performed significantly worse than healthy controls on the MorGen, driven by differences in adjective performance. When controlling for aphasia severity, those with fluent aphasia performed significantly better in their production of nouns, plurals, number, size, and color than those who had non-fluent aphasia, but both groups were similarly inclined to omit genitive marking. Aim 3: Lesions in the territory of the temporal branch of the posterior cerebral artery were associated with poorer performance in nouns, size, and color. Lesions in the territory of the anterior cerebral artery were associated with poorer performance in numbers. Conclusions: This work highlights the value of the MorGen as a tool for post-stroke language evaluation that complements the skills captured in more widely-used assessments such as the WAB and BNT.

Lesion-symptom Mapping of Acceptability Judgments in Chronic Poststroke Aphasia Reveals the Neurobiological Underpinnings of Receptive Syntax.

Disagreements persist regarding the neural basis of syntactic processing, which has been linked both to inferior frontal and posterior temporal regions of the brain. One focal point of the debate concerns the role of inferior frontal areas in receptive syntactic ability, which is mostly assessed using sentence comprehension involving complex syntactic structures, a task that is potentially confounded with working memory. Syntactic acceptability judgments may provide a better measure of receptive syntax by reducing the need to use high working memory load and complex sentences and by enabling assessment of various types of syntactic violations. We therefore tested the perception of grammatical violations by people with poststroke aphasia (n = 25), along with matched controls (n = 16), using English sentences involving errors in word order, agreement, or subcategorization. Lesion data were also collected. Control participants performed near ceiling in accuracy with higher discriminability of agreement and subcategorization violations than word order; aphasia participants were less able to discriminate violations, but, on average, paralleled control participants discriminability of types of violations. Lesion-symptom mapping showed a correlation between discriminability and posterior temporal regions, but not inferior frontal regions. We argue that these results diverge from models holding that frontal areas are amodal core regions in syntactic structure building and favor models that posit a core hierarchical system in posterior temporal regions.

Grammatical Parallelism in Aphasia: A Lesion-Symptom Mapping Study.

Sentence structure, or syntax, is potentially a uniquely creative aspect of the human mind. Neuropsychological experiments in the 1970s suggested parallel syntactic production and comprehension deficits in agrammatic Broca's aphasia, thought to result from damage to syntactic mechanisms in Broca's area in the left frontal lobe. This hypothesis was sometimes termed overarching agrammatism, converging with developments in linguistic theory concerning central syntactic mechanisms supporting language production and comprehension. However, the evidence supporting an association among receptive syntactic deficits, expressive agrammatism, and damage to frontal cortex is equivocal. In addition, the relationship among a distinct grammatical production deficit in aphasia, paragrammatism, and receptive syntax has not been assessed. We used lesion-symptom mapping in three partially overlapping groups of left-hemisphere stroke patients to investigate these issues: grammatical production deficits in a primary group of 53 subjects and syntactic comprehension in larger sample sizes (N = 130, 218) that overlapped with the primary group. Paragrammatic production deficits were significantly associated with multiple analyses of syntactic comprehension, particularly when incorporating lesion volume as a covariate, but agrammatic production deficits were not. The lesion correlates of impaired performance of syntactic comprehension were significantly associated with damage to temporal lobe regions, which were also implicated in paragrammatism, but not with the inferior and middle frontal regions implicated in expressive agrammatism. Our results provide strong evidence against the overarching agrammatism hypothesis. By contrast, our results suggest the possibility of an alternative grammatical parallelism hypothesis rooted in paragrammatism and a central syntactic system in the posterior temporal lobe.

Lexico-semantics obscures lexical syntax.

A recently emerging generalization about language and the brain is that brain regions implicated in language that show syntax-related activations (e.g., increased activation for more complex sentence structures) also tend to show word-related activations, such as increased activation to reading real words (e.g. RAIN) relative to pseudowords (e.g. PHREZ) (Fedorenko et al., 2016). Fedorenko et al., (2020) generalize as follows: "…syntactic/combinatorial processing is not separable from lexico-semantic processing at the level of brain regions-or even voxel subsets-within the language network". Based on this generalization, Fedorenko et al. have made the conclusion "…that a cognitive architecture whereby syntactic processing is not separable from the processing of individual word meanings is most likely", arguing against "syntax-centric" views of language as promulgated by Chomsky and others. However, the notion of "lexico-semantics" articulated here obscures the fact that words are both syntactic and semantic entities. Because of this, any functional neuroimaging experiment that manipulates lexicality will almost assuredly tax both syntactic and semantic resources, and is therefore inadequate for isolating conceptual-semantic processing in the brain in exclusion to syntax. In addition, Fedorenko et al. do not satisfactorily account for robust lesion data that shows clear functional-anatomical dissociations within the language network. Finally, a "syntax-centric" view of language is perfectly compatible with Fedorenko et al.'s conclusions about syntax-selectivity in the brain because of the multiple potential mappings between linguistic theory and neurobiology beyond individual brain regions.

The Wernicke conundrum revisited: evidence from connectome-based lesion-symptom mapping.

Wernicke's area has been assumed since the 1800s to be the primary region supporting word and sentence comprehension. However, in 2015 and 2019, Mesulam and colleagues raised what they termed the 'Wernicke conundrum', noting widespread variability in the anatomical definition of this area and presenting data from primary progressive aphasia that challenged this classical assumption. To resolve the conundrum, they posited a 'double disconnection' hypothesis: that word and sentence comprehension deficits in stroke-based aphasia result from disconnection of anterior temporal and inferior frontal regions from other parts of the brain due to white matter damage, rather than dysfunction of Wernicke's area itself. To test this hypothesis, we performed lesion-deficit correlations, including connectome-based lesion-symptom mapping, in four large, partially overlapping groups of English-speaking chronic left hemisphere stroke survivors. After removing variance due to object recognition and associative semantic processing, the same middle and posterior temporal lobe regions were implicated in both word comprehension deficits and complex non-canonical sentence comprehension deficits. Connectome lesion-symptom mapping revealed similar temporal-occipital white matter disconnections for impaired word and non-canonical sentence comprehension, including the temporal pole. We found an additional significant temporal-parietal disconnection for non-canonical sentence comprehension deficits, which may indicate a role for phonological working memory in processing complex syntax, but no significant frontal disconnections. Moreover, damage to these middle-posterior temporal lobe regions was associated with both word and non-canonical sentence comprehension deficits even when accounting for variance due to the strongest anterior temporal and inferior frontal white matter disconnections, respectively. Our results largely agree with the classical notion that Wernicke's area, defined here as middle superior temporal gyrus and middle-posterior superior temporal sulcus, supports both word and sentence comprehension, suggest a supporting role for temporal pole in both word and sentence comprehension, and speak against the hypothesis that comprehension deficits in Wernicke's aphasia result from double disconnection.

The Cortical Organization of Syntactic Processing Is Supramodal: Evidence from American Sign Language.

Areas within the left-lateralized neural network for language have been found to be sensitive to syntactic complexity in spoken and written language. Previous research has revealed that these areas are active for sign language as well, but whether these areas are specifically responsive to syntactic complexity in sign language independent of lexical processing has yet to be found. To investigate the question, we used fMRI to neuroimage deaf native signers' comprehension of 180 sign strings in American Sign Language (ASL) with a picture-probe recognition task. The ASL strings were all six signs in length but varied at three levels of syntactic complexity: sign lists, two-word sentences, and complex sentences. Syntactic complexity significantly affected comprehension and memory, both behaviorally and neurally, by facilitating accuracy and response time on the picture-probe recognition task and eliciting a left lateralized activation response pattern in anterior and posterior superior temporal sulcus (aSTS and pSTS). Minimal or absent syntactic structure reduced picture-probe recognition and elicited activation in bilateral pSTS and occipital-temporal cortex. These results provide evidence from a sign language, ASL, that the combinatorial processing of anterior STS and pSTS is supramodal in nature. The results further suggest that the neurolinguistic processing of ASL is characterized by overlapping and separable neural systems for syntactic and lexical processing.

Functional differentiation in the language network revealed by lesion-symptom mapping.

Theories of language organization in the brain commonly posit that different regions underlie distinct linguistic mechanisms. However, such theories have been criticized on the grounds that many neuroimaging studies of language processing find similar effects across regions. Moreover, condition by region interaction effects, which provide the strongest evidence of functional differentiation between regions, have rarely been offered in support of these theories. Here we address this by using lesion-symptom mapping in three large, partially-overlapping groups of aphasia patients with left hemisphere brain damage due to stroke (N = 121, N = 92, N = 218). We identified multiple measure by region interaction effects, associating damage to the posterior middle temporal gyrus with syntactic comprehension deficits, damage to posterior inferior frontal gyrus with expressive agrammatism, and damage to inferior angular gyrus with semantic category word fluency deficits. Our results are inconsistent with recent hypotheses that regions of the language network are undifferentiated with respect to high-level linguistic processing.

One cat, Two cats, Red cat, Blue cats: Eliciting morphemes from individuals with primary progressive aphasia.

BACKGROUND: Progressive neurodegenerative impairment with central language features, primary progressive aphasia (PPA), can be further distinguished for many individuals into one of three variants: semantic, non-fluent/agrammatic, and logopenic variant PPA. Variants differ in their relative preservation and deficits of language skills, particularly in word finding and grammar. The majority of elicited language assessments used in this population focus on single noun and verb production, while modifiers and inflectional morphemes are far less commonly examined. AIMS: The purpose of the present study is to determine whether there was an interaction between PPA variant and production of high-frequency nouns, proper names, modifiers, and bound inflectional morphemes to better understand how the variants differ. METHODS & PROCEDURES: Forty-six people with PPA and 47 individuals with no known neurological diagnoses completed a morphosyntactic generation assessment designed to target differential production of high-frequency nouns, proper names, modifiers (number, size, color), and bound inflectional morphemes (plural -s and possessive 's), the Morphosyntactic Generation test (MorGen). Performance is averaged for each of the seven morphosyntactic targets independently, resulting in seven separate performance scores. OUTCOMES & RESULTS: Individuals with PPA performed significantly more poorly than controls on the assessed morphemes in a repeated-measures analysis of variance, as well as on each morpheme considered independently via t-test.In a multivariable analysis of variance among PPA variants, the interaction of morpheme and PPA variant was significant, suggesting different variants produced the morphemes with a significantly different pattern of success. When morphemes were considered independently, only production of colour resulted in a significant difference between variants, driven by the performance of individuals with nfavPPA, who performed near-ceiling. When MorGen performance was used to predict PPA variant in a multinomial logistic regression the model was significant, with age, plural -s, noun, and number contributing significantly to the prediction. In a discriminate function analysis, classification of cases was best for agrammatic variant with 70% accuracy. CONCLUSIONS: Individuals with PPA, particularly semantic and logopenic variants, demonstrated difficulty on the MorGen compared to controls. The MorGen proved useful in predicting PPA variant. These findings highlight the potential benefit of examining a broader range of morphemes, particularly bound morphemes and modifiers, in addition to the more frequently investigated classes of nouns and verbs when understanding PPA.

A double dissociation between plural and possessive "s": Evidence from the Morphosyntactic Generation test.

People with aphasia demonstrate impaired production of bound inflectional morphemes, such as noun plurals and possession. They often show greater difficulty in marking possession versus plurality. Using a new tool for eliciting language, the Morphosyntactic Generation test, we assessed people with primary progressive aphasia and those in the acute and chronic phase following left hemisphere stroke. Clinical profiles were associated with different strengths and weaknesses in language production. Performance of the plural was stronger than possessive in group analyses. However, some individuals demonstrated the inverse pattern of performance. These participants provide counter-evidence to the theory that difficulty with marking possessives is purely the result of their greater cognitive-linguistic complexity and support a functional double dissociation between possessives and plurals. The deficits resulted from morphosyntactic impairment. Future work is needed to understand why plural and possessive markers were differently sensitive to neurological disorders of language.

The Cortical Organization of Syntax.

Syntax, the structure of sentences, enables humans to express an infinite range of meanings through finite means. The neurobiology of syntax has been intensely studied but with little consensus. Two main candidate regions have been identified: the posterior inferior frontal gyrus (pIFG) and the posterior middle temporal gyrus (pMTG). Integrating research in linguistics, psycholinguistics, and neuroscience, we propose a neuroanatomical framework for syntax that attributes distinct syntactic computations to these regions in a unified model. The key theoretical advances are adopting a modern lexicalized view of syntax in which the lexicon and syntactic rules are intertwined, and recognizing a computational asymmetry in the role of syntax during comprehension and production. Our model postulates a hierarchical lexical-syntactic function to the pMTG, which interconnects previously identified speech perception and conceptual-semantic systems in the temporal and inferior parietal lobes, crucial for both sentence production and comprehension. These relational hierarchies are transformed via the pIFG into morpho-syntactic sequences, primarily tied to production. We show how this architecture provides a better account of the full range of data and is consistent with recent proposals regarding the organization of phonological processes in the brain.

Syntax-Sensitive Regions of the Posterior Inferior Frontal Gyrus and the Posterior Temporal Lobe Are Differentially Recruited by Production and Perception.

Matchin and Hickok (2020) proposed that the left posterior inferior frontal gyrus (PIFG) and the left posterior temporal lobe (PTL) both play a role in syntactic processing, broadly construed, attributing distinct functions to these regions with respect to production and perception. Consistent with this hypothesis, functional dissociations between these regions have been demonstrated with respect to lesion-symptom mapping in aphasia. However, neuroimaging studies of syntactic comprehension typically show similar activations in these regions. In order to identify whether these regions show distinct activation patterns with respect to syntactic perception and production, we performed an fMRI study contrasting the subvocal articulation and perception of structured jabberwocky phrases (syntactic), sequences of real words (lexical), and sequences of pseudowords (phonological). We defined two sets of language-selective regions of interest (ROIs) in individual subjects for the PIFG and the PTL using the contrasts [syntactic > lexical] and [syntactic > phonological]. We found robust significant interactions of comprehension and production between these 2 regions at the syntactic level, for both sets of language-selective ROIs. This suggests a core difference in the function of these regions with respect to production and perception, consistent with the lesion literature.

Agrammatism and Paragrammatism: A Cortical Double Dissociation Revealed by Lesion-Symptom Mapping.

The fundamental distinction of grammatical deficits in aphasia, agrammatism and paragrammatism, was made over a century ago. However, the extent to which the agrammatism/paragrammatism distinction exists independently of differences in speech fluency has not clearly been investigated. Despite much research on agrammatism, the lesion correlates of paragrammatism are essentially unknown. Lesion-symptom mapping was used to investigate the degree to which the lesion correlates of agrammatism and paragrammatism overlap or dissociate. Four expert raters assessed videos of 53 right-handed patients with aphasia following chronic left-hemisphere stroke retelling the Cinderella story. Consensus discussion determined each subject's classification with respect to grammatical deficits as Agrammatic, Paragrammatic, Both, or No Grammatical Deficit. Each subject's lesion was manually drawn on a high-resolution MRI and warped to standard space for group analyses. Lesion-symptom mapping analyses were performed in NiiStat including lesion volume as a covariate. Secondary analyses included speech rate (words per minute) as an additional covariate. Region of interest analyses identified a double dissociation between these syndromes: damage to Broca's area was significantly associated with agrammatism, p = 0.001 (but not paragrammatism, p = 0.930), while damage to the left posterior superior and middle temporal gyri was significantly associated with paragrammatism, p < 0.001 (but not agrammatism, p = 0.873). The same results obtained when regressing out the effect of speech rate, and nonoverlapping lesion distributions between the syndromes were confirmed by uncorrected whole brain analyses. Our results support a fundamental distinction between agrammatism and paragrammatism.

Same words, different structures: An fMRI investigation of argument relations and the angular gyrus.

In fMRI, increased activation for combinatorial syntactic and semantic processing is typically observed in a set of left hemisphere brain areas: the angular gyrus (AG), the anterior temporal lobe (ATL), the posterior superior temporal sulcus (pSTS), and the inferior frontal gyrus (IFG). Recent work has suggested that semantic combination is supported by the ATL and the AG, with a division of labor in which AG is involved in event concepts and ATL is involved in encoding conceptual features of entities and/or more general forms of semantic combination. The current fMRI study was designed to refine hypotheses about the angular gyrus processes in question. In particular, we ask whether the AG supports the computation of argument structure (a linguistic notion that depends on a verb taking other phrases as arguments) or the computation of event concepts more broadly. To distinguish these possibilities we used a novel, lexically-matched contrast: noun phrases (NP) (the frightened boy) and verb phrases (VP) (frightened the boy), where VPs contained argument structure, denoting an event and assigning a thematic role to its argument, and NPs did not, denoting only a semantically enriched entity. Results showed that while many regions showed increased activity for NPs and VPs relative to unstructured word lists (AG, ATL, pSTS, anterior IFG), replicating evidence of their involvement in combinatorial processing, neither AG or ATL showed differences in activation between the VP and NP conditions. These results suggest that increased AG activity does not reflect the computation of argument structure per se, but are compatible with a view in which the AG represents event information denoted by words such as frightened independent of their grammatical context. By contrast, pSTS and posterior IFG did show increased activation for the VPs relative to NPs. We suggest that these effects may reflect differences in syntactic processing and working memory engaged by different structural relations.

The temporal dynamics of structure and content in sentence comprehension: Evidence from fMRI-constrained MEG.

Humans have a striking capacity to combine words into sentences that express new meanings. Previous research has identified key brain regions involved in this capacity, but little is known about the time course of activity in these regions, as hemodynamic methods such as fMRI provide little insight into temporal dynamics of neural activation. We performed an MEG experiment to elucidate the temporal dynamics of structure and content processing within four brain regions implicated by fMRI data from the same experiment: the temporo-parietal junction (TPJ), the posterior temporal lobe (PTL), the anterior temporal lobe (ATL), and the anterior inferior frontal gyrus (IFG). The TPJ showed increased activity for both structure and content near the end of the sentence, consistent with a role in incremental interpretation of event semantics. The PTL, a region not often associated with core aspects of syntax, showed a strong early effect of structure, consistent with predictive parsing models, and both structural and semantic context effects on function words. These results provide converging evidence that the PTL plays an important role in lexicalized syntactic processing. The ATL and IFG, regions traditionally associated with syntax, showed minimal effects of sentence structure. The ATL, PTL and IFG all showed effects of semantic content: increased activation for real words relative to nonwords. Our fMRI-guided MEG investigation therefore helps identify syntactic and semantic aspects of sentence comprehension in the brain in both spatial and temporal dimensions.

Phonological Feature Repetition Suppression in the Left Inferior Frontal Gyrus.

Models of speech production posit a role for the motor system, predominantly the posterior inferior frontal gyrus, in encoding complex phonological representations for speech production, at the phonemic, syllable, and word levels [Roelofs, A. A dorsal-pathway account of aphasic language production: The WEAVER++/ARC model. Cortex, 59(Suppl. C), 33-48, 2014; Hickok, G. Computational neuroanatomy of speech production. Nature Reviews Neuroscience, 13, 135-145, 2012; Guenther, F. H. Cortical interactions underlying the production of speech sounds. Journal of Communication Disorders, 39, 350-365, 2006]. However, phonological theory posits subphonemic units of representation, namely phonological features [Chomsky, N., & Halle, M. The sound pattern of English, 1968; Jakobson, R., Fant, G., & Halle, M. Preliminaries to speech analysis. The distinctive features and their correlates. Cambridge, MA: MIT Press, 1951], that specify independent articulatory parameters of speech sounds, such as place and manner of articulation. Therefore, motor brain systems may also incorporate phonological features into speech production planning units. Here, we add support for such a role with an fMRI experiment of word sequence production using a phonemic similarity manipulation. We adapted and modified the experimental paradigm of Oppenheim and Dell [Oppenheim, G. M., & Dell, G. S. Inner speech slips exhibit lexical bias, but not the phonemic similarity effect. Cognition, 106, 528-537, 2008; Oppenheim, G. M., & Dell, G. S. Motor movement matters: The flexible abstractness of inner speech. Memory & Cognition, 38, 1147-1160, 2010]. Participants silently articulated words cued by sequential visual presentation that varied in degree of phonological feature overlap in consonant onset position: high overlap (two shared phonological features; e.g., /r/ and /l/) or low overlap (one shared phonological feature, e.g., /r/ and /b/). We found a significant repetition suppression effect in the left posterior inferior frontal gyrus, with increased activation for phonologically dissimilar words compared with similar words. These results suggest that phonemes, particularly phonological features, are part of the planning units of the motor speech system.

Neural networks supporting audiovisual integration for speech: A large-scale lesion study.

Auditory and visual speech information are often strongly integrated resulting in perceptual enhancements for audiovisual (AV) speech over audio alone and sometimes yielding compelling illusory fusion percepts when AV cues are mismatched, the McGurk-MacDonald effect. Previous research has identified three candidate regions thought to be critical for AV speech integration: the posterior superior temporal sulcus (STS), early auditory cortex, and the posterior inferior frontal gyrus. We assess the causal involvement of these regions (and others) in the first large-scale (N = 100) lesion-based study of AV speech integration. Two primary findings emerged. First, behavioral performance and lesion maps for AV enhancement and illusory fusion measures indicate that classic metrics of AV speech integration are not necessarily measuring the same process. Second, lesions involving superior temporal auditory, lateral occipital visual, and multisensory zones in the STS are the most disruptive to AV speech integration. Further, when AV speech integration fails, the nature of the failure-auditory vs visual capture-can be predicted from the location of the lesions. These findings show that AV speech processing is supported by unimodal auditory and visual cortices as well as multimodal regions such as the STS at their boundary. Motor related frontal regions do not appear to play a role in AV speech integration.

A neuronal retuning hypothesis of sentence-specificity in Broca's area.

It is clear that the left inferior frontal gyrus (LIFG) contributes in some fashion to sentence processing. While neuroimaging and neuropsychological evidence support a domain-general working memory function, recent neuroimaging data show that particular subregions of the LIFG, particularly the pars triangularis (pTri), show selective activation for sentences relative to verbal working memory and cognitive control tasks. These data suggest a language-specific function rather than a domain-general one. To resolve this apparent conflict, I propose separating claims of domain-generality and specificity independently for computations and representations-a given brain region may respond to a specific representation while performing a general computation over that representation, one shared with other systems. I hypothesize that the pTri underlies a language-specific working memory system, comprised of general memory retrieval/attention operations specialized for syntactic representations. There is a parallelism of top-down retrieval function among the phonological and semantic levels, localized to the pars opercularis and pars orbitalis, respectively. I further explore the idea of how such a system emerges in the human brain through the framework of neuronal retuning: the "borrowing" of domain-general mechanisms for language, either in evolution or development. The empirical data appear to tentatively support a developmental account of language-specificity in the pTri, possibly through connections to the posterior superior temporal sulcus (pSTS), a region that is both anatomically distinct for humans and functionally essential for language. Evidence of representational response specificity obtained from neuroimaging studies is useful in understanding how cognition is implemented in the brain. However, understanding the shared computations across domains and neural systems is necessary for a fuller understanding of this problem, providing potential answers to questions of how specialized systems, such as language, are implemented in the brain.

Neural evidence for predictive coding in auditory cortex during speech production.

Recent models of speech production suggest that motor commands generate forward predictions of the auditory consequences of those commands, that these forward predications can be used to monitor and correct speech output, and that this system is hierarchically organized (Hickok, Houde, & Rong, Neuron, 69(3), 407--422, 2011; Pickering & Garrod, Behavior and Brain Sciences, 36(4), 329--347, 2013). Recent psycholinguistic research has shown that internally generated speech (i.e., imagined speech) produces different types of errors than does overt speech (Oppenheim & Dell, Cognition, 106(1), 528--537, 2008; Oppenheim & Dell, Memory & Cognition, 38(8), 1147-1160, 2010). These studies suggest that articulated speech might involve predictive coding at additional levels than imagined speech. The current fMRI experiment investigates neural evidence of predictive coding in speech production. Twenty-four participants from UC Irvine were recruited for the study. Participants were scanned while they were visually presented with a sequence of words that they reproduced in sync with a visual metronome. On each trial, they were cued to either silently articulate the sequence or to imagine the sequence without overt articulation. As expected, silent articulation and imagined speech both engaged a left hemisphere network previously implicated in speech production. A contrast of silent articulation with imagined speech revealed greater activation for articulated speech in inferior frontal cortex, premotor cortex and the insula in the left hemisphere, consistent with greater articulatory load. Although both conditions were silent, this contrast also produced significantly greater activation in auditory cortex in dorsal superior temporal gyrus in both hemispheres. We suggest that these activations reflect forward predictions arising from additional levels of the perceptual/motor hierarchy that are involved in monitoring the intended speech output.

The role of the IFG and pSTS in syntactic prediction: Evidence from a parametric study of hierarchical structure in fMRI.

Sentences encode hierarchical structural relations among words. Several neuroimaging experiments aiming to localize combinatory operations responsible for creating this structure during sentence comprehension have contrasted short, simple phrases and sentences to unstructured controls. Some of these experiments have revealed activation in the left inferior frontal gyrus (IFG) and posterior superior temporal sulcus (pSTS), associating these regions with basic syntactic combination. However, the wide variability of these effects across studies raises questions about this interpretation. In an fMRI experiment, we provide support for an alternative hypothesis: these regions underlie top-down syntactic predictions that facilitate sentence processing but are not necessary for building syntactic structure. We presented stimuli with three levels of structure: unstructured lists, two-word phrases, and simple, short sentences; and two levels of content: natural stimuli with real words and stimuli with open-class items replaced with pseudowords (jabberwocky). While both the phrase and sentence conditions engaged syntactic combination, our experiment only encouraged syntactic prediction in the sentence condition. We found increased activity for both natural and jabberwocky sentences in the left IFG (pars triangularis and pars orbitalis) and pSTS relative to unstructured word lists and two-word phrases, but we did not find any such effects for two-word phrases relative to unstructured word lists in these areas. Our results are most consistent with the hypothesis that increased activity in IFG and pSTS for basic contrasts of structure reflects syntactic prediction. The pars opercularis of the IFG showed a response profile consistent with verbal working memory. We found incremental effects of structure in the anterior temporal lobe (ATL), and increased activation only for sentences in the angular gyrus (AG)/temporal-parietal junction (TPJ) - both regions showed these effects for stimuli with all real words. These findings support a role for the ATL in semantic combination and the AG/TPJ in thematic processing.