Friedemann Pulvermüller

Active perception: sensorimotor circuits as a cortical basis for language

Action and perception are functionally linked in the brain, but a hotly debated question is whether perception and comprehension of stimuli depend on motor circuits. Brain language mechanisms are ideal for addressing this question. Neuroimaging investigations have found specific motor activations when subjects understand speech sounds, word meanings and sentence structures. Moreover, studies involving transcranial magnetic stimulation and patients with lesions affecting inferior frontal regions of the brain have shown contributions of motor circuits to the comprehension of phonemes, semantic categories and grammar. These data show that language comprehension benefits from frontocentral action systems, indicating that action and perception circuits are interdependent.

Functional links between motor and language systems

Transcranial magnetic stimulation (TMS) was applied to motor areas in the left language‐dominant hemisphere while right‐handed human subjects made lexical decisions on words related to actions. Response times to words referring to leg actions (e.g. kick) were compared with those to words referring to movements involving the arms and hands (e.g. pick). TMS of hand and leg areas influenced the processing of arm and leg words differentially, as documented by a significant interaction of the factors Stimulation site and Word category. Arm area TMS led to faster arm than leg word responses and the reverse effect, faster lexical decisions on leg than arm words, was present when TMS was applied to leg areas. TMS‐related differences between word categories were not seen in control conditions, when TMS was applied to hand and leg areas in the right hemisphere and during sham stimulation. Our results show that the left hemispheric cortical systems for language and action are linked to each other in a category‐specific manner and that activation in motor and premotor areas can influence the processing of specific kinds of words semantically related to arm or leg actions. By demonstrating specific functional links between action and language systems during lexical processing, these results call into question modular theories of language and motor functions and provide evidence that the two systems interact in the processing of meaningful information about language and action.

Constraint-Induced Therapy of Chronic Aphasia After Stroke

Patients with chronic aphasia were assigned randomly to a group to receive either conventional aphasia therapy or constraint-induced (CI) aphasia therapy, a new therapeutic technique requiring intense practice over a relatively short period of consecutive days. CI aphasia therapy is realized in a communicative therapeutic environment constraining patients to practice systematically speech acts with which they have difficulty. Patients in both groups received the same amount of treatment (30 to 35 hours) as 10 days of massed-practice language exercises for the CI aphasia therapy group (3 hours per day minimum; 10 patients) or over a longer period of ≈4 weeks for the conventional therapy group (7 patients). CI aphasia therapy led to significant and pronounced improvements on several standard clinical tests, on self-ratings, and on blinded-observer ratings of the patients’ communicative effectiveness in everyday life. Patients who received the control intervention failed to achieve comparable improvements. Data suggest that the language skills of patients with chronic aphasia can be improved in a short period by use of an appropriate massed-practice technique that focuses on the patients’ communicative needs.

Motor cortex maps articulatory features of speech sounds

The processing of spoken language has been attributed to areas in the superior temporal lobe, where speech stimuli elicit the greatest activation. However, neurobiological and psycholinguistic models have long postulated that knowledge about the articulatory features of individual phonemes has an important role in their perception and in speech comprehension. To probe the possible involvement of specific motor circuits in the speech-perception process, we used event-related functional MRI and presented experimental subjects with spoken syllables, including [p] and [t] sounds, which are produced by movements of the lips or tongue, respectively. Physically similar nonlinguistic signal-correlated noise patterns were used as control stimuli. In localizer experiments, subjects had to silently articulate the same syllables and, in a second task, move their lips or tongue. Speech perception most strongly activated superior temporal cortex. Crucially, however, distinct motor regions in the precentral gyrus sparked by articulatory movements of the lips and tongue were also differentially activated in a somatotopic manner when subjects listened to the lip- or tongue-related phonemes. This sound-related somatotopic activation in precentral gyrus shows that, during speech perception, specific motor circuits are recruited that reflect phonetic distinctive features of the speech sounds encountered, thus providing direct neuroimaging support for specific links between the phonological mechanisms for speech perception and production.

The time course of visual word recognition as revealed by linear regression analysis of ERP data

EEG correlates of a range of psycholinguistic word properties were used to investigate the time course of access to psycholinguistic information during visual word recognition. Neurophysiological responses recorded in a visual lexical decision task were submitted to linear regression analysis. First, 10 psycholinguistic features of each of 300 stimulus words were submitted to a principal component analysis, which yielded four orthogonal variables likely to reflect separable processes in visual word recognition: Word length, Letter n-gram frequency, Lexical frequency and Semantic coherence of a words morphological family. Since the lexical decision task required subjects to distinguish between words and pseudowords, the binary variable Lexicality was also investigated using a factorial design. Word-pseudoword differences in the event-related potential first appeared at 160 ms after word onset. However, regression analysis of EEG data documented a much earlier effect of both Word length and Letter n-gram frequency around 90 ms. Lexical frequency showed its earliest effect slightly later, at 110 ms, and Semantic coherence significantly correlated with neurophysiological measures around 160 ms, simultaneously with the lexicality effect. Source estimates indicated parieto-temporo-occipital generators for the factors Length, Letter n-gram frequency and Word frequency, but widespread activation with foci in left anterior temporal lobe and inferior frontal cortex related to Semantic coherence. At later stages (>200 ms), all variables exhibited simultaneous EEG correlates. These results indicate that information about surface form and meaning of a lexical item is first accessed at different times in different brain systems and then processed simultaneously, thus supporting cascaded interactive processing models.

Brain reflections of words and their meaning

The neurobiological organization of meaningful language units, morphemes and words, has been illuminated by recent metabolic and neurophysiological imaging studies. When humans process words from different categories, sets of cortical areas become active differentially. The meaning of a word, more precisely aspects of its reference, may be crucial for determining which set of cortical areas is involved in its processing. Word-related neuron webs with specific cortical distributions might underlie the observed category-specific differences in brain activity. Neuroscientific principles can explain these differential topographies.

Conceptual representations in mind and brain: Theoretical developments, current evidence and future directions

Conceptual representations in long-term memory crucially contribute to perception and action, language and thought. However, the precise nature of these conceptual memory traces is discussed controversially. In particular, the grounding of concepts in the sensory and motor brain systems is the focus of a current debate. Here, we review theoretical accounts of the structure and neural basis of conceptual memory and evaluate them in light of recent empirical evidence. Models of conceptual processing can be distinguished along four dimensions: (i) amodal versus modality-specific, (ii) localist versus distributed, (iii) innate versus experience-dependent, and (iv) stable versus flexible. A systematic review of behavioral and neuroimaging studies in healthy participants along with brain-damaged patients will then be used to evaluate the competing theoretical approaches to conceptual representations. These findings indicate that concepts are flexible, distributed representations comprised of modality-specific conceptual features. Conceptual features are stored in distinct sensory and motor brain areas depending on specific sensory and motor experiences during concept acquisition. Three important controversial issues are highlighted, which require further clarification in future research: the existence of an amodal conceptual representation in the anterior temporal lobe, the causal role of sensory and motor activation for conceptual processing and the grounding of abstract concepts in perception and action. We argue that an embodiment view of conceptual representations realized as distributed sensory and motor cell assemblies that are complemented by supramodal integration brain circuits may serve as a theoretical framework to guide future research on concrete and abstract concepts.

Grasping Ideas with the Motor System: Semantic Somatotopy in Idiom Comprehension

Single words and sentences referring to bodily actions activate the motor cortex. However, this semantic grounding of concrete language does not address the critical question whether the sensory–motor system contributes to the processing of abstract meaning and thought. We examined functional magnetic resonance imaging activation to idioms and literal sentences including arm- and leg-related action words. A common left fronto-temporal network was engaged in sentence reading, with idioms yielding relatively stronger activity in (pre)frontal and middle temporal cortex. Crucially, somatotopic activation along the motor strip, in central and precentral cortex, was elicited by idiomatic and literal sentences, reflecting the body part reference of the words embedded in the sentences. Semantic somatotopy was most pronounced after sentence ending, thus reflecting sentence-level processing rather than that of single words. These results indicate that semantic representations grounded in the sensory–motor system play a role in the composition of sentence-level meaning, even in the case of idioms.

Effects of word length and frequency on the human event-related potential

OBJECTIVE We investigated the influence of the length and frequency of printed words on the amplitude and peak latencies of event-related potentials (ERPs). This served two goals, namely (I) to clarify their possible effects as confounds in ERP experiments employing word-stimuli, and (II) to determine the point in time of lexical access in visual word recognition. METHODS EEG was recorded from 64 scalp sites while subjects (n=12) performed a lexical decision task. Word length and frequency were orthogonally varied between stimulus groups, whereas variables including regularity of spelling and orthographic tri-gram frequency were kept constant. RESULTS Long words produced the strongest brain response early on (approximately 100 ms after stimulus onset), whereas those to short words became strongest later (150-360 ms). Lower ERP amplitudes were elicited by words with high frequency compared with low frequency words in the latency ranges 150-190 ms and 320-360 ms. However, we did not find evidence for a robust alteration of peak latencies with word frequency. CONCLUSIONS Length and frequency of word stimuli have independent and additive effects on the amplitude of the ERP. Studies on the precise time course of cognitive processes should consider their potentially confounding character. Our data support the view that lexical access takes place as early as 150 ms after onset of written word stimuli.

Memory traces for words as revealed by the Mismatch Negativity

Brain responses to the same spoken syllable completing a Finnish word or a pseudo-word were studied. Native Finnish-speaking subjects were instructed to ignore the sound stimuli and watch a silent movie while the mismatch negativity (MMN), an automatic index of experience-dependent auditory memory traces, was recorded. The MMN to each syllable was larger when it completed a word than when it completed a pseudo-word. This enhancement, reaching its maximum amplitude at about 150 ms after the words recognition point, did not occur in foreign subjects who did not know any Finnish. These results provide the first demonstration of the presence of memory traces for individual spoken words in the human brain. Using whole-head magnetoencephalography, the major intracranial source of this word-related MMN was found in the left superior temporal lobe.