Papers

2017

The extent to which language processing involves prediction of upcoming inputs remains a question of ongoing debate. One important data point comes from DeLong et al. (2005) who reported that an N400-like event-related potential correlated with a probabilistic index of upcoming input. This result is often cited as evidence for gradient probabilistic prediction of form and/or semantics, prior to the bottom-up input becoming available. However, a recent multi-lab study reports a failure to find these effects (Nieuwland et al., 2017). We review the evidence from both studies, including differences in the design and analysis approach between them. Building on over a decade of research on prediction since DeLong et al. (2005)’s original study, we also begin to spell out the computational nature of predictive processes that one might expect to correlate with ERPs that are evoked by a functional element whose form is dependent on an upcoming predicted word. For paradigms with this type of design, we propose an index of anticipatory processing, Bayesian surprise, and apply it to the updating of semantic predictions. We motivate this index both theoretically and empirically. We show that, for studies of the type discussed here, Bayesian surprise can be closely approximated by another, more easily estimated information theoretic index, the surprisal (or Shannon information) of the input. We re-analyze the data from Nieuwland and colleagues using surprisal rather than raw probabilities as an index of prediction. We find that surprisal is gradiently correlated with the amplitude of the N400, even in the data shared by Nieuwland and colleagues. Taken together, our review suggests that the evidence from both studies is compatible with anticipatory semantic processing. We do, however, emphasize the need for future studies to further clarify the nature and degree of form prediction, as well as its neural signatures, during language comprehension.

2016

Weber, K., Lau, E. F., Stillerman, B., & Kuperberg, G. R. (2016). The Yin and the Yang of Prediction: An fMRI Study of Semantic Predictive Processing. PLoS One, 11(3), e0148637. https://doi.org/10.1371/journal.pone.0148637

Probabilistic prediction plays a crucial role in language comprehension. When predictions are fulfilled, the resulting facilitation allows for fast, efficient processing of ambiguous, rapidly-unfolding input; when predictions are not fulfilled, the resulting error signal allows us to adapt to broader statistical changes in this input. We used functional Magnetic Resonance Imaging to examine the neuroanatomical networks engaged in semantic predictive processing and adaptation. We used a relatedness proportion semantic priming paradigm, in which we manipulated the probability of predictions while holding local semantic context constant. Under conditions of higher (versus lower) predictive validity, we replicate previous observations of reduced activity to semantically predictable words in the left anterior superior/middle temporal cortex, reflecting facilitated processing of targets that are consistent with prior semantic predictions. In addition, under conditions of higher (versus lower) predictive validity we observed significant differences in the effects of semantic relatedness within the left inferior frontal gyrus and the posterior portion of the left superior/middle temporal gyrus. We suggest that together these two regions mediated the suppression of unfulfilled semantic predictions and lexico-semantic processing of unrelated targets that were inconsistent with these predictions. Moreover, under conditions of higher (versus lower) predictive validity, a functional connectivity analysis showed that the left inferior frontal and left posterior superior/middle temporal gyrus were more tightly interconnected with one another, as well as with the left anterior cingulate cortex. The left anterior cingulate cortex was, in turn, more tightly connected to superior lateral frontal cortices and subcortical regions-a network that mediates rapid learning and adaptation and that may have played a role in switching to a more predictive mode of processing in response to the statistical structure of the wider environmental context. Together, these findings highlight close links between the networks mediating semantic prediction, executive function and learning, giving new insights into how our brains are able to flexibly adapt to our environment.

Fields, E. C., & Kuperberg, G. R. (2016). Dynamic Effects of Self-Relevance and Task on the Neural Processing of Emotional Words in Context. Frontiers in Psychology, 6, 2003. https://doi.org/10.3389/fpsyg.2015.02003

We used event-related potentials (ERPs) to examine the interactions between task, emotion, and contextual self-relevance on processing words in social vignettes. Participants read scenarios that were in either third person (other-relevant) or second person (self-relevant) and we recorded ERPs to a neutral, pleasant, or unpleasant critical word. In a previously reported study (Fields and Kuperberg, 2012) with these stimuli, participants were tasked with producing a third sentence continuing the scenario. We observed a larger LPC to emotional words than neutral words in both the self-relevant and other-relevant scenarios, but this effect was smaller in the self-relevant scenarios because the LPC was larger on the neutral words (i.e., a larger LPC to self-relevant than other-relevant neutral words). In the present work, participants simply answered comprehension questions that did not refer to the emotional aspects of the scenario. Here we observed quite a different pattern of interaction between self-relevance and emotion: the LPC was larger to emotional vs. neutral words in the self-relevant scenarios only, and there was no effect of self-relevance on neutral words. Taken together, these findings suggest that the LPC reflects a dynamic interaction between specific task demands, the emotional properties of a stimulus, and contextual self-relevance. We conclude by discussing implications and future directions for a functional theory of the emotional LPC.

Kuperberg, G. R. (2016). Separate streams or probabilistic inference? What the N400 can tell us about the comprehension of events. Language, Cognition and Neuroscience, 31(5), 602-616. https://doi.org/10.1080/23273798.2015.1130233

Since the early 2000s, several ERP studies have challenged the assumption that we always use syntactic contextual information to influence semantic processing of incoming words, as reflected by the N400 component. One approach for explaining these findings is to posit distinct semantic and syntactic processing mechanisms, each with distinct time courses. While this approach can explain specific datasets, it cannot account for the wider body of findings. I propose an alternative explanation: a dynamic generative framework in which our goal is to infer the underlying event that best explains the set of inputs encountered at any given time. Within this framework, combinations of semantic and syntactic cues with varying reliabilities are used as evidence to weight probabilistic hypotheses about this event. I further argue that the computational principles of this framework can be extended to understand how we infer situation models during discourse comprehension, and intended messages during spoken communication.

Kuperberg, G. R., & Jaeger, F. (2016). What do we mean by prediction in language comprehension?. Language, Cognition and Neuroscience, 31(1), 32-59. https://doi.org/10.1080/23273798.2015.1102299

We consider several key aspects of prediction in language comprehension: its computational nature, the representational level(s) at which we predict, whether we use higher level representations to predictively pre-activate lower level representations, and whether we ’commit’ in any way to our predictions, beyond pre-activation. We argue that the bulk of behavioral and neural evidence suggests that we predict probabilistically and at multiple levels and grains of representation. We also argue that we can, in principle, use higher level inferences to predictively pre-activate information at multiple lower representational levels. We also suggest that the degree and level of predictive pre-activation might be a function of the expected utility of prediction, which, in turn, may depend on comprehenders’ goals and their estimates of the relative reliability of their prior knowledge and the bottom-up input. Finally, we argue that all these properties of language understanding can be naturally explained and productively explored within a multi-representational hierarchical actively generative architecture whose goal is to infer the message intended by the producer, and in which predictions play a crucial role in explaining the bottom-up input.

Delaney-Busch, N., Wilkie, G., & Kuperberg, G. R. (2016). Vivid: How valence and arousal influence word processing under different task demands. Cognitive, Affective, & Behavioral Neuroscience, 16(3), 413-432.

In this study, we used event-related potentials to examine how different dimensions of emotion—valence and arousal—influence different stages of word processing under different task demands. In two experiments, two groups of participants viewed the same single emotional and neutral words while carrying out different tasks. In both experiments, valence (pleasant, unpleasant, and neutral) was fully crossed with arousal (high and low). We found that the task made a substantial contribution to how valence and arousal modulated the late positive complex (LPC), which is thought to reflect sustained evaluative processing (particularly of emotional stimuli). When participants performed a semantic categorization task in which emotion was not directly relevant to task performance, the LPC showed a larger amplitude for high-arousal than for low-arousal words, but no effect of valence. In contrast, when participants performed an overt valence categorization task, the LPC showed a large effect of valence (with unpleasant words eliciting the largest positivity), but no effect of arousal. These data show not only that valence and arousal act independently to influence word processing, but that their relative contributions to prolonged evaluative neural processes are strongly influenced by the situational demands (and by individual differences, as revealed in a subsequent analysis of subjective judgments).

Lau, E. F., Weber, K., Gramfort, A., Hämäläinen, M. S., & Kuperberg, G. R. (2016). Spatiotemporal Signatures of Lexical-Semantic Prediction. Cerebral Cortex, 26(4), 1377-87. https://doi.org/10.1093/cercor/bhu219 (Original work published 2016)

Although there is broad agreement that top-down expectations can facilitate lexical-semantic processing, the mechanisms driving these effects are still unclear. In particular, while previous electroencephalography (EEG) research has demonstrated a reduction in the N400 response to words in a supportive context, it is often challenging to dissociate facilitation due to bottom-up spreading activation from facilitation due to top-down expectations. The goal of the current study was to specifically determine the cortical areas associated with facilitation due to top-down prediction, using magnetoencephalography (MEG) recordings supplemented by EEG and functional magnetic resonance imaging (fMRI) in a semantic priming paradigm. In order to modulate expectation processes while holding context constant, we manipulated the proportion of related pairs across 2 blocks (10 and 50% related). Event-related potential results demonstrated a larger N400 reduction when a related word was predicted, and MEG source localization of activity in this time-window (350-450 ms) localized the differential responses to left anterior temporal cortex. fMRI data from the same participants support the MEG localization, showing contextual facilitation in left anterior superior temporal gyrus for the high expectation block only. Together, these results provide strong evidence that facilitatory effects of lexical-semantic prediction on the electrophysiological response 350-450 ms postonset reflect modulation of activity in left anterior temporal cortex.

2015

Language and thought dysfunction are central to the schizophrenia syndrome. They are evident in the major symptoms of psychosis itself, particularly as disorganized language output (positive thought disorder) and auditory verbal hallucinations (AVHs), and they also manifest as abnormalities in both high-level semantic and contextual processing and low-level perception. However, the literatures characterizing these abnormalities have largely been separate and have sometimes provided mutually exclusive accounts of aberrant language in schizophrenia. In this review, we propose that recent generative probabilistic frameworks of language processing can provide crucial insights that link these four lines of research. We first outline neural and cognitive evidence that real-time language comprehension and production normally involve internal generative circuits that propagate probabilistic predictions to perceptual cortices - predictions that are incrementally updated based on prediction error signals as new inputs are encountered. We then explain how disruptions to these circuits may compromise communicative abilities in schizophrenia by reducing the efficiency and robustness of both high-level language processing and low-level speech perception. We also argue that such disruptions may contribute to the phenomenology of thought-disordered speech and false perceptual inferences in the language system (i.e., AVHs). This perspective suggests a number of productive avenues for future research that may elucidate not only the mechanisms of language abnormalities in schizophrenia, but also promising directions for cognitive rehabilitation.

Fogel, A. R., Rosenberg, J. C., Lehman, F. M., Kuperberg, G. R., & Patel, A. D. (2015). Studying Musical and Linguistic Prediction in Comparable Ways: The Melodic Cloze Probability Method. Frontiers in Psychology, 6, 1718. https://doi.org/10.3389/fpsyg.2015.01718

Prediction or expectancy is thought to play an important role in both music and language processing. However, prediction is currently studied independently in the two domains, limiting research on relations between predictive mechanisms in music and language. One limitation is a difference in how expectancy is quantified. In language, expectancy is typically measured using the cloze probability task, in which listeners are asked to complete a sentence fragment with the first word that comes to mind. In contrast, previous production-based studies of melodic expectancy have asked participants to sing continuations following only one to two notes. We have developed a melodic cloze probability task in which listeners are presented with the beginning of a novel tonal melody (5-9 notes) and are asked to sing the note they expect to come next. Half of the melodies had an underlying harmonic structure designed to constrain expectations for the next note, based on an implied authentic cadence (AC) within the melody. Each such ’authentic cadence’ melody was matched to a ’non-cadential’ (NC) melody matched in terms of length, rhythm and melodic contour, but differing in implied harmonic structure. Participants showed much greater consistency in the notes sung following AC vs. NC melodies on average. However, significant variation in degree of consistency was observed within both AC and NC melodies. Analysis of individual melodies suggests that pitch prediction in tonal melodies depends on the interplay of local factors just prior to the target note (e.g., local pitch interval patterns) and larger-scale structural relationships (e.g., melodic patterns and implied harmonic structure). We illustrate how the melodic cloze method can be used to test a computational model of melodic expectation. Future uses for the method include exploring the interplay of different factors shaping melodic expectation, and designing experiments that compare the cognitive mechanisms of prediction in music and language.

Takaya, S., Kuperberg, G. R., Liu, H., Greve, D. N., Makris, N., & Stufflebeam, S. M. (2015). Asymmetric projections of the arcuate fasciculus to the temporal cortex underlie lateralized language function in the human brain. Frontiers in Neuroanatomy, 9, 119. https://doi.org/10.3389/fnana.2015.00119

The arcuate fasciculus (AF) in the human brain has asymmetric structural properties. However, the topographic organization of the asymmetric AF projections to the cortex and its relevance to cortical function remain unclear. Here we mapped the posterior projections of the human AF in the inferior parietal and lateral temporal cortices using surface-based structural connectivity analysis based on diffusion MRI and investigated their hemispheric differences. We then performed the cross-modal comparison with functional connectivity based on resting-state functional MRI (fMRI) and task-related cortical activation based on fMRI using a semantic classification task of single words. Structural connectivity analysis showed that the left AF connecting to Broca’s area predominantly projected in the lateral temporal cortex extending from the posterior superior temporal gyrus to the mid part of the superior temporal sulcus and the middle temporal gyrus, whereas the right AF connecting to the right homolog of Broca’s area predominantly projected to the inferior parietal cortex extending from the mid part of the supramarginal gyrus to the anterior part of the angular gyrus. The left-lateralized projection regions of the AF in the left temporal cortex had asymmetric functional connectivity with Broca’s area, indicating structure-function concordance through the AF. During the language task, left-lateralized cortical activation was observed. Among them, the brain responses in the temporal cortex and Broca’s area that were connected through the left-lateralized AF pathway were specifically correlated across subjects. These results suggest that the human left AF, which structurally and functionally connects the mid temporal cortex and Broca’s area in asymmetrical fashion, coordinates the cortical activity in these remote cortices during a semantic decision task. The unique feature of the left AF is discussed in the context of the human capacity for language.

See also: Structural MRI