Peter Indefrey

The spatial and temporal signatures of word production components

This paper presents the results of a comprehensive meta-analysis of the relevant imaging literature on word production (82 experiments). In addition to the spatial overlap of activated regions, we also analyzed the available data on the time course of activations. The analysis specified regions and time windows of activation for the core processes of word production: lexical selection, phonological code retrieval, syllabification, and phonetic/articulatory preparation. A comparison of the word production results with studies on auditory word/non-word perception and reading showed that the time course of activations in word production is, on the whole, compatible with the temporal constraints that perception processes impose on the production processes they affect in picture/word interference paradigms.

The Spatial and Temporal Signatures of Word Production Components: A Critical Update

In the first decade of neurocognitive word production research the predominant approach was brain mapping, i.e., investigating the regional cerebral brain activation patterns correlated with word production tasks, such as picture naming and word generation. Indefrey and Levelt (2004) conducted a comprehensive meta-analysis of word production studies that used this approach and combined the resulting spatial information on neural correlates of component processes of word production with information on the time course of word production provided by behavioral and electromagnetic studies. In recent years, neurocognitive word production research has seen a major change toward a hypothesis-testing approach. This approach is characterized by the design of experimental variables modulating single component processes of word production and testing for predicted effects on spatial or temporal neurocognitive signatures of these components. This change was accompanied by the development of a broader spectrum of measurement and analysis techniques. The article reviews the findings of recent studies using the new approach. The time course assumptions of Indefrey and Levelt (2004) have largely been confirmed requiring only minor adaptations. Adaptations of the brain structure/function relationships proposed by Indefrey and Levelt (2004) include the precise role of subregions of the left inferior frontal gyrus as well as a probable, yet to date unclear role of the inferior parietal cortex in word production.

The Neural Circuitry Involved in the Reading of German Words and Pseudowords: A PET Study

Silent reading and reading aloud of German words and pseudowords were used in a PET study using (15O) butanol to examine the neural correlates of reading and of the phonological conversion of legal letter strings, with or without meaning. The results of 11 healthy, right-handed volunteers in the age range of 25 to 30 years showed activation of the lingual gyri during silent reading in comparison with viewing a fixation cross. Comparisons between the reading of words and pseudo-words suggest the involvement of the middle temporal gyri in retrieving both the phonological and semantic code for words. The reading of pseudowords activates the left inferior frontal gyrus, including the ventral part of Brocas area, to a larger extent than the reading of words. This suggests that this area might be involved in the sublexical conversion of orthographic input strings into phonological output codes. (Pre)motor areas were found to be activated during both silent reading and reading aloud. On the basis of the obtained activation patterns, it is hypothesized that the articulation of high-frequency syllables requires the retrieval of their concomitant articulatory gestures from the SMA and that the articulation of low-frequency syllables recruits the left medial premotor cortex.

Syntactic processing in left prefrontal cortex is independent of lexical meaning.

In language comprehension a syntactic representation is built up even when the input is semantically uninterpretable. We report data on brain activation during syntactic processing, from an experiment on the detection of grammatical errors in meaningless sentences. The experimental paradigm was such that the syntactic processing was distinguished from other cognitive and linguistic functions. The data reveal that in syntactic error detection an area of the left dorsolateral prefrontal cortex, adjacent to Brocas area, is specifically involved in the syntactic processing aspects, whereas other prefrontal areas subserve general error detection processes.

The Neurobiology of Language Beyond Single Words

A hallmark of human language is that we combine lexical building blocks retrieved from memory in endless new ways. This combinatorial aspect of language is referred to as unification. Here we focus on the neurobiological infrastructure for syntactic and semantic unification. Unification is characterized by a high-speed temporal profile including both prediction and integration of retrieved lexical elements. A meta-analysis of numerous neuroimaging studies reveals a clear dorsal/ventral gradient in both left inferior frontal cortex and left posterior temporal cortex, with dorsal foci for syntactic processing and ventral foci for semantic processing. In addition to core areas for unification, further networks need to be recruited to realize language-driven communication to its full extent. One example is the theory of mind network, which allows listeners and readers to infer the intended message (speaker meaning) from the coded meaning of the linguistic utterance. This indicates that sensorimotor simulation cannot handle all of language processing.

Reading in a Regular Orthography: An fMRI Study Investigating the Role of Visual Familiarity

In order to separate the cognitive processes associated with phonological encoding and the use of a visual word form lexicon in reading, it is desirable to compare the processing of words presented in a visually familiar form with words in a visually unfamiliar form. Japanese Kana orthography offers this possibility. Two phonologically equivalent but visually dissimilar syllabaries allow the writing of, for example, foreign loanwords in two ways, only one of which is visually familiar. Familiarly written words, unfamiliarly written words, and pseudowords were presented in both Kana syllabaries (yielding six conditions in total) to participants during an fMRI measurement with a silent articulation task (Experiment 1) and a phonological lexical decision task (Experiment 2) using an event-related design. Consistent over two experimental tasks, the three different stimulus types (familiar, unfamiliar, and pseudoword) were found to activate selectively different brain regions previously associated with phonological encoding and word retrieval or meaning. Compatible with the predictions of the dual-route model for reading, pseudowords and visually unfamiliar words, which have to be read using phonological assembly, caused an increase in brain activity in left inferior frontal regions (BA 44/47), as compared to visually familiar words. Visually familiar and unfamiliar words were found to activate a range of areas associated with lexico-semantic processing more strongly than pseudowords, such as the left and right temporo-parietal region (BA 39/40), a region in the left middle/inferior temporal gyrus (BA 20/21), and the posterior cingulate (BA 31).

Neural Responses to the Production and Comprehension of Syntax in Identical Utterances.

Following up on an earlier positron emission tomography (PET) experiment (Indefrey et al., 2001), we used a scene description paradigm to investigate whether a posterior inferior frontal region subserving syntactic encoding for speaking is also involved in syntactic parsing during listening. In the language production part of the experiment, subjects described visually presented scenes using either sentences, sequences of noun phrases, or sequences of syntactically unrelated words. In the language comprehension part of the experiment, subjects were auditorily presented with the same kinds of utterances and judged whether they matched the visual scenes. We were able to replicate the previous finding of a region in caudal Brocas area that is sensitive to the complexity of syntactic encoding in language production. In language comprehension, no hemodynamic activation differences due to syntactic complexity were found. Given that correct performance in the judgment task did not require syntactic processing of the auditory stimuli, the results suggest that the degree to which listeners recruit syntactic processing resources in language comprehension may be a function of the syntactic demands of the task or the stimulus material.

Equivalent responses to lexical and nonlexical visual stimuli in occipital cortex: a functional magnetic resonance imaging study

Stimulus-related changes in cerebral blood oxygenation were measured using high-resolution functional magnetic resonance imaging sequentially covering visual occipital areas in contiguous sections. During dynamic imaging, healthy subjects silently viewed pseudowords, single false fonts, or length-matched strings of the same false fonts. The paradigm consisted of a sixfold alternation of an activation and a control task. With pseudowords as activation vs single false fonts as control, responses were seen mainly in medial occipital cortex. These responses disappeared when pseudowords were alternated with false font strings as the control and reappeared when false font strings instead of pseudowords served as activation and were alternated with single false fonts. The string-length contrast alone, therefore, is sufficient to account for the activation pattern observed in medial visual cortex when word-like stimuli are contrasted with single characters.

An inverse relation between event-related and time-frequency violation responses in sentence processing.

The relationship between semantic and grammatical processing in sentence comprehension was investigated by examining event-related potential (ERP) and event-related power changes in response to semantic and grammatical violations. Sentences with semantic, phrase structure, or number violations and matched controls were presented serially (1.25 words/s) to 20 participants while EEG was recorded. Semantic violations were associated with an N400 effect and a theta band increase in power, while grammatical violations were associated with a P600 effect and an alpha/beta band decrease in power. A quartile analysis showed that for both types of violations, larger average violation effects were associated with lower relative amplitudes of oscillatory activity, implying an inverse relation between ERP amplitude and event-related power magnitude change in sentence processing.

Syntactic priming in German–English bilinguals during sentence comprehension

A longstanding question in bilingualism is whether syntactic information is shared between the two language processing systems. We used an fMRI repetition suppression paradigm to investigate syntactic priming in reading comprehension in German-English late-acquisition bilinguals. In comparison to conventional subtraction analyses in bilingual experiments, repetition suppression has the advantage of being able to detect neuronal populations that are sensitive to properties that are shared by consecutive stimuli. In this study, we manipulated the syntactic structure between prime and target sentences. A sentence with a passive sentence structure in English was preceded either by a passive or by an active sentence in English or German. We looked for repetition suppression effects in left inferior frontal, left precentral and left middle temporal regions of interest. These regions were defined by a contrast of all non-target sentences in German and English versus the baseline of sentence-format consonant strings. We found decreases in activity (repetition suppression effects) in these regions of interest following the repetition of syntactic structure from the first to the second language and within the second language. Moreover, a separate behavioural experiment using a word-by-word reading paradigm similar to the fMRI experiment showed faster reading times for primed compared to unprimed English target sentences regardless of whether they were preceded by an English or a German sentence of the same structure. We conclude that there is interaction between the language processing systems and that at least some syntactic information is shared between a bilinguals languages with similar syntactic structures.