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MRC CBU project SL4
Brain Dynamics Of Language In Space And Time
Theory guided research on the brain basis of language has been undertaken using MEG, EEG, fMRI, TMS and behavioural experiments in healthy subjects and language-impaired patients. Subproject SL4.1/2 focussed on the interplay between language systems in the brain and action and perception systems. SL4.3 focussed on the time course of language-related brain processes using a new tool for studying language processing, the cognitive Mismatch Negativity. Hemispheric interaction in language processing and language therapy were, respectively, highlighted in SL4.4 and SL4.5.
The neurobiological model of language (NML, Pulvermüller, 1999) predicts that the neuroanatomical links between cortical areas and the mapping of correlated neuronal activity onto connection strength yield the development of distributed cortical circuits for processing language elements. These distributed circuits bind, at the neurofunctional level, neurons in different areas of cortex with critical roles in action and perception together with additional neurons in "higher", non-sensory and non-motor, areas. The circuits carry specific phonological, lexical, morphological, syntactic and semantic functions immanent to language. The neuronal circuits are distributed over different sets of cortical areas and activated according to different time schedules. Cortical distribution of the circuits, in part, reflects their cognitive/linguistic function (Pulvermüller, 2005; Pulvermüller & Shtyrov, 2006; Pulvermüller, 2008b).
We investigated the neural basis of linguistic processes at a more fine grained level of specificity than most common studies of language processes, where general "phonological networks", the "lexicon", the "semantic system" or "grammar" are typically targeted. In contrast, we focussed on a finer grain size of specificity: phonemes of different kinds were contrasted with each other, as were also lexical items with different morphosyntactic and semantic features; in syntax, we targeted specific constructions that are of theoretical interest (agreement). Our results revealed brain mechanisms reflecting such fine-grained distinctions, some of which confirmed predictions of the NML.
SL4.1 and 4.2 Language and the brain's action and perception systems
Semantic somatotopy/topography
In the domain of semantics, we could show that word meaning can be mapped on cortical activations at a level of detail not previously attained. Fine-grained semantic categories of words matched in relevant physical and psycholinguistic parameters elicited differential activation patterns in the brain, which was revealed by multimodal imaging techniques.
The critical prediction of the neurobiological model of language had been that the language and action systems of the human brain are not encapsulated modules, but rather highly interactive systems and that these systems are co-activated when words of specific semantic kinds are being processed. The semantic somatotopy model (SSM, Pulvermüller, 1999; 2001) predicts that the meaning of action words is reflected by activation in the motor strip of the human brain. We found that indeed words that refer to actions usually involving specific body parts – for example lick, pick and kick, which, respectively, imply face, arm and leg actions – activate the parts of motor and premotor cortex that control the respective actions (Figure SL4.1, Pulvermüller et al., 2000; Hauk et al., 2004; Hauk & Pulvermüller, 2004b; Shtyrov et al., 2004; Pulvermüller et al., 2005a; Pulvermüller et al., 2005c; Hauk et al., 2008a; Hauk et al., 2008b; Pulvermüller, 2008b; 2008a). The result that action language sparks semantically related areas of motor systems has also been replicated externally (e.g., Tettamanti et al., 2005; Aziz-Zadeh et al., 2006; Kemmerer et al., 2008).
Words referring to objects primarily known through the visual modality produce more activation in specific parts of the ventral visual stream for object processing compared with control words (Chao et al., 1999; Pulvermüller et al., 1999). We added further, more detailed results that also words definable primarily on the basis of different types of visual information produce different activation patterns. Words semantically related to colour and to shape (red vs. round) activated different inferior-temporal areas (in the fusiform and parahippocampal gyri, Moscoso Del Prado Martin et al., 2006; Pulvermüller & Hauk, 2006). A further point has been made about words implying odours, which were found to specifically activate the olfactory system (González et al., 2006). Together with the action word findings, these results document a much finer grain of semantic distinctions in the brain than some previous cognitive theories assumed (cf. animal vs. tool and living vs. nonliving distinctions, see also Humphreys & Forde, 2001; Martin, 2007; Barsalou, 2008). Furthermore, our results rule out earlier attempts to attribute category differences in brain activation to confounding psycholinguistic features, imageability, grammatical complexity or the semantic similarity between items (e.g., Bird et al., 2000; Tyler et al., 2000), as these properties were controlled in our studies.
Phonological somatotopy
Speech sounds may be based on different brain circuits in temporal cortex (Braitenberg & Pulvermüller, 1992; Diesch et al., 1996; Obleser et al., 2006) and in the motor system. In the same way as semantic features of action words can be mapped onto the motor system of the brain, speech sounds, phonemes, do relate to specific areas of the motor cortex because they are produced with specific articulators. For instance, the tongue contributes strongly to the production of a [t] whereas a [p] draws more heavily on the lips. Investigating this issue with fMRI, we found that listening to speech sounds activates the specific motor areas contributing to the production of a perceived sound (Pulvermüller et al., 2006a). In a similar manner, listening to non-linguistic click sounds produced with fingers or tongue activates the respective motor regions (Hauk et al., 2006c), a fact reminiscent of mirror neuron activation to sounds in the motor cortex (Kohler et al., 2002). It will be important in future to define differences in the cortical activation patterns reflecting semantic and phonological knowledge linked to speech and written language.
It had been remarked that the pure activation of sensorimotor systems in the brain does not prove a contribution at the functional level to semantics, phonology, or any other linguistic or cognitive function. We have argued that, to support a role of a brain area in a specific linguistic/cognitive process (e.g. action semantics), it also needs to be shown that the brain area is (i) instantaneously activated, (ii) activated across different tasks and (iii) necessary for the process. To address (i) and to rule out late "epiphenomenal" activation, we have used EEG/MEG methods and found semantic somatotopy effects to be near-instantaneous (delays of <200ms upon delivery of stimulus information, Pulvermüller et al., 2001a; Hauk & Pulvermüller, 2004b; Shtyrov et al., 2004; Pulvermüller et al., 2005c; Hauk et al., 2008b). Regarding (ii), we could confirm semantic somatotopy in a range of task, for example lexical decision, passive silent reading and even under distraction, both to speech and to written language recorded in both EEG and MEG (Pulvermüller et al., 2001a; Hauk & Pulvermüller, 2004b; Shtyrov et al., 2004; Pulvermüller et al., 2005c). The necessity of motor cortex for lexicosemantic processing is indicated by a range of patient studies (e.g., Bak et al., 2001; Kemmerer & Tranel, 2003; Neininger & Pulvermüller, 2003; Bak et al., 2006; Cotelli et al., 2006; Boulenger et al., 2008). We also used TMS and found differential influence of arm/leg motor cortex stimulation on arm-/leg-word processing (Pulvermüller et al., 2005a). Although more work is needed to confirm the level of category specificity documented by imaging results in clinical populations and to further explore the range of tasks that might or might not lead to category effects (see Future proposal SL4), the results so far leave little doubt about the close and specific functional links between motor and language cortices (Pulvermüller, 2005; , 2008b).
SL4.3 Psycholinguistic information access in brain-space and –time
A widely held view had been that psycholinguistic information is accessed serially, in a stepwise fashion, starting with access to phonological and syntactic information about lexical items, followed by lexicosemantic and morphological information access and finally syntactic in-depth processing (Friederici, 2002). A range of both serial and cascaded psycholinguistic models view lexical access and semantics as relatively late processes in language comprehension (e.g., Dell et al., 1997; Norris et al., 2000), whose earliest manifestation might be in the N400 component of the event-related brain response (Kutas & Federmeier, 2000; Kiyonaga et al., 2007). As psycholinguistic work had indicated early access to different kinds of psycholinguistic information (e.g., Marslen-Wilson & Tyler, 1975), we investigated the time course of psycholinguistic information access in the written and spoken modality using MEG and EEG and found consistent evidence for early (latency 100-200ms) near simultaneous neural indexes of phonological/orthographic, lexical, syntactic, morphological and semantic information immanent to words and sentences. This research used both conventional factorial designs crossing relevant variables (while matching others) and multiple regression designs.
In the written modality, word length and word frequency were found to have early (<200ms) neurophysiological correlates in both MEG (Assadollahi & Pulvermüller, 2003) and EEG (Hauk & Pulvermüller, 2004a), with indication for early interaction effects revealed by MEG only. As word length can reflect processes at different levels (orthographic, physical etc.), letter bi- and trigram frequency was contrasted with lexicality in subsequent work suggesting an early but slightly different (100 vs. 160ms) appearance of the sublexical and lexical effects (Hauk et al., 2006b). Parallel results were found for processing object pictures (Hauk et al., 2007), suggesting that more general processing levels of feature and object/word representations are indexed by the physiological activity at 100 and 160ms.
Differences in brain activation between single words can also be investigated in multiple regression designs. Whereas exact matching of stimuli and orthogonal factorial designs can prove the involvement of a brain area in a few selected cognitive/linguistic processes (e.g., Fiez et al., 1999), regression designs have the advantage that the effect of multiple variables can be studied in one experiment while stimulus matching (which sometimes implies the choice of atypical items) is not necessary. When applying a multi-regression design, we confirmed an early EEG manifestation of bi/trigram frequencies of words (<100ms), followed by word length and word frequency (120ms) and lexicality together with semantics (160ms) (Hauk et al., 2006a). As word length is confounded with the number N of neighbours of a written word, a subsequent study showed independent effects of these two variables shortly after 100ms. Our results generally suggest that visual word processing is aligned in time with that of other kinds of familiar visual stimuli, including objects and faces, with a critical activation time for representations at the object level at ~150-200ms (Bentin et al., 1996; Zion-Golumbic & Bentin, 2007).
It has been argued that lexical and lexicosemantic information may be accessed first but that the integration of a word into its semantic context would certainly need more time, probably ~400ms (Barber & Kutas, 2007). On the other hand, we have argued that comparatively large stimulus variance could underlie the absence of early effects in a range of experiments (Pulvermüller, 1999; Pulvermüller & Shtyrov, 2006). This was tested using an N400 paradigm with orthogonal variation of word length, word frequency and the cloze probability with which words appear in a given sentence context. The results revealed main effects of cloze probability and word frequency only at >300 ms, whereas early-on (100-200 ms) interactions of these variables with word lengths predominated. This proves that early effects also exist for semantic context integration and, critically, that increasing stimulus variance (here word length) abolishes these early effects (Penolazzi et al., 2007; Pulvermüller, 2007).
Speech processing was studied using a variation of the paradigm established to record the Mismatch Negativity (MMN) brain response. We chose this paradigm, because the MMN (1) reflects the activation of linguistic memory traces in the brain (Näätänen et al., 1997; Pulvermüller et al., 2001b), (2) makes it possible to record neurophysiological activity to single well-controlled stimuli thus reducing stimulus variance to a minimum, and (3) can be obtained in the absence of focussed attention to the eliciting stimuli, removing confounds of attention and stimulus-specific strategy variations. Importantly, the MMN design makes it possible to fully orthogonalize stimulus properties (e.g. composite sounds) and the critical variable (e.g., lexicality: for example in the quadruplet pipe, pite*, bipe*, bite; real words in italics, pseudowords marked with *). If effects appear in orthogonalised MMN designs, the conclusions are justified that they survive the attenuating effect of stimulus repetition (Dale et al., 2000), are not due to physical stimulus features or their variance, and are to a degree independent of attention being focussed on them (Pulvermüller & Shtyrov, 2006; Shtyrov & Pulvermüller, 2007b). General results are summarized in Figure 4.2.
Using the MMN, we found early MEG and EEG manifestations of lexical features, within 200 ms after the word recognition point (WRP, Marslen-Wilson, 1987). Earlier work on spoken language had shown that the language comprehension system engages in comprehension and semantic processing on the basis of incomplete information (Müller & Kutas, 1996; Van Petten et al., 1999; Hagoort & Brown, 2000). We found that words produce larger MMNs than pseudowords around 130-150ms after the WRP where words are first recognized with confidence (Pulvermüller et al., 2001b; Shtyrov & Pulvermüller, 2002; Endrass et al., 2004). Differential activation to unattended words and pseudowords was confirmed by fMRI (Shtyrov et al., 2008). Critically, we found that WRP latencies correlate with the latency of a temporal source of the MMN (Pulvermüller et al., 2006b). Lexicality effects in the MMN paradigm were replicated by other labs (e.g., Pettigrew et al., 2004).
In the same early time range (~150ms after WRP), brain responses to the same chirp-like acoustic stimulus indicate whether it is perceived as a noise, a speech sound or as a grammatical affix (Shtyrov et al., 2005). Semantic word properties are also reflected in the MMN at 100-200 ms (Pulvermüller et al., 2004), with different semantic types eliciting semantic MMN effects at different latencies (Shtyrov et al., 2004; Pulvermüller et al., 2005c). Leg-action words activate the dorsal frontocentral cortex with a slight delay (30ms) upon the inferiorfrontal activation seen to face- and arm-action words, a difference consistent with cortico-cortical conduction delays.
Effects of syntactic violations, which had been reported early and late in ERP work on spoken language (Friederici et al., 1993; Hagoort et al., 1993), surfaced in MMN paradigms at early latencies (~150ms upon WRP). This was found in a range of languages (English, German, Finnish) using MEG and EEG (Pulvermüller & Shtyrov, 2003; Shtyrov et al., 2003; Pulvermüller & Shtyrov, 2006). Interestingly, the early effects were found to reflect grammaticality rather than sequential probability (Pulvermüller & Assadollahi, 2007), which constitutes neurophysiological support for discrete theories of grammar processing and the rule concept (Pinker, 1994). The grammatical constructions eliciting the syntactic MMN included number/case/gender agreement between noun and verb and between determiner and noun, thus arguing that grammar mechanisms drawing outside and above the range of classic phrase structure rules are processed early-on. We have developed a theoretical model of brain circuits for grammar and syntax that takes account of these findings (Pulvermüller, 2003b; 2003a; Knoblauch & Pulvermüller, 2005). Results on the syntactic MMN have been replicated by other labs (Menning et al., 2005; Hasting et al., 2007). The automaticity of the syntactic MMN effect at early latencies (<150ms) has been proven by a recent study (Pulvermüller et al., 2008).
Semantic context integration effects in the MMN paradigm were also recently reported (Shtyrov & Pulvermüller, 2007a). Not surprisingly, these were found as early (<150 ms) as in previous reports on semantic and pragmatic violations (e.g., Brown et al., 2000) and in the visual modality (Sereno et al., 2003).
These psychophysiological results on written and spoken language processing provide consistent evidence for an early time course of psycholinguistic information access, with all information types being accessed near-simultaneously, within 100-200ms after critical stimulus information comes in, therefore confirming conclusions from behavioural data (Marslen-Wilson & Tyler, 1975). Differences in activation delays between information types (e.g., sublexical vs. lexical, semantic subcategories) are explained by conduction delays between cortical areas (10-50 ms, Pulvermüller, 2003a). We recently started a new project on explaining cortical activation spreading on the basis of the NML implemented in a neuroanatomically realistic neurocomputational model of the language cortex (Wennekers et al., 2006; Garagnani et al., 2007; 2008).
SL4.4 Laterality and hemispheric interaction
Language is left-lateralised in most right-handers but some critical contribution of the right hemisphere to language processing is also evident (Zaidel, 1976; Pulvermüller & Mohr, 1996). One paradigm used to argue in favour of bihemispheric interactive contributions to language processing is the redundant bilateral stimulation paradigm with the same information sent to the left or right hemisphere, or, in a third bilateral-redundant condition, simultaneously to both (Mohr et al., 1994). Experiments using this paradigm show that lexical information sent simultaneously to both hemispheres leads to better performance on lexical decision tasks than unilateral stimulation of the dominant hemisphere. We were able to record for the first time neurophysiological indexes of the bilateral redundancy gain to spoken and written words (Endrass et al., 2004; Mohr et al., 2007). Source localisation indicates that activity in left-hemispheric language areas is enhanced in the bilateral condition for words but not pseudowords. This was found in different tasks, including lexical decision and the unattended oddball task. Clinical populations with functional deficits of the corpus callosum – especially split brain patients but also patients with schizophrenia – did not show the bilateral redundancy gain for words at the behavioural or physiological level (Mohr et al., 2008). This pattern of results confirms earlier predictions of a model of interhemispheric interaction in lexical processing (Pulvermüller & Mohr, 1996).
SL4.5 Translational language research: Aphasia therapy
Whereas classical wisdom had been that aphasia does usually not improve significantly when the chronic stage is reached 1 year after onset of the disease (Pedersen et al., 1995), we developed a new intensive language-action therapy (ILAT), also called constraint-induced aphasia therapy, and showed, in a randomized controlled trial, that its intensive application over a short time period (two weeks) led to significant improvement of language performance in patients with chronic aphasia (Pulvermüller et al., 2001c).
The key for success of the new method lies in the rigorous application of knowledge available from basic research in neuroscience. Brain research documented that the cortical mechanisms for language and action are tightly interwoven (see SL4.1/2) and, concurrently, the new approach to language therapy in neurological patients implements language training in the context of relevant linguistic and non-linguistic actions, therefore taking advantage of the mutual connections of language and action systems in the brain. A further well-known neuroscience principle is that learning at the neuronal level is driven by correlation; consequently, the new approach to language therapy emphasizes massed practice in a short time, thus maximizing therapy quantity and frequency and, therefore, correlation at the behavioural and neuronal levels. Learned non-use of unsuccessful actions plays a major role in the chronification of neurological deficits and the behavioural approach to therapy has therefore employed shaping and other learning techniques to counteract such non-use (cf. Taub et al., 2002).
We studied the effect of intensive language-action therapy on brain activation and found bilateral increase of cortical activation to words over the two-week period during which therapy improved performance on clinical language tests and tests of everyday communications. There was a significant correlation between left- and right-hemispheric source strengths and the improvement on clinical tests, confirming the role of both hemispheres in language generally (SL4.4, see also Pulvermüller et al., 2004) and in language recovery specifically (Pulvermüller et al., 2005b). In a recent review, we discuss perspectives for further improving speech-language therapy, including drug treatment that may be particularly fruitful when applied in conjunction with language-action therapy (Pulvermüller & Berthier, 2008). Pulvermüller collaborates with Prof. Berthier's group at the Centre of Medical and Health Research (University of Malaga) to optimize ILAT methods (supported by Pfizer and Lundbek).
The newly developed intensive language-action therapy is an efficient tool for improving language functions even at chronic stages of aphasia. Therapy studies using this rapid technique to improve language functions can open new perspectives for research into the plasticity of human language circuits. Intensive language-action therapy constitutes a milestone in documenting the fruitful interaction between neuroscience research at the theoretical level (on language-action connections) and its clinical translational application.
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