Experiential grounding of abstract concepts: Processing of abstract mental state concepts engages brain regions involved in mentalizing, automatic speech, and lip movements.

concepts like mental state concepts lack a physical referent, which can be directly perceived. Classical theories therefore claim that abstract concepts require amodal representations detached from experiential brain systems. However, grounded cognition approaches suggest an involvement of modal experiential brain regions in the processing of abstract concepts. In the present functional magnetic resonance imaging study, we investigated the relation of the processing of abstract mental state concepts to modal experiential brain systems in a fine-grained fashion. Participants performed lexical decisions on abstract mental state as well as on verbal association concepts as control category. Experiential brain systems related to the processing of mental states, generating verbal associations, automatic speech as well as hand and lip movements were determined by corresponding localizer tasks. Processing of abstract mental state concepts neuroanatomically overlapped with activity patterns associated with processing of mental states, generating verbal associations, automatic speech and lip movements. Hence, mental state concepts activate the mentalizing brain network, complemented by perceptual-motor brain regions involved in simulation of visual or action features associated with social interactions, linguistic brain regions as well as face-motor brain regions recruited for articulation. The present results provide compelling evidence for the rich grounding of abstract mental state concepts in experiential brain systems related to mentalizing, verbal communication and mouth action.

Academic training increases grounding of scientific concepts in experiential brain systems.

Scientific concepts typically transcendent our sensory experiences. Traditional approaches to science education therefore assume a shift towards amodal or verbal knowledge representations during academic training. Grounded cognition approaches, in contrast, predict a maintenance of grounding of the concepts in experiential brain networks or even an increase. To test these competing approaches, the present study investigated the semantic content of scientific psychological concepts and identified the corresponding neural circuits using functional magnetic resonance imaging (fMRI) in undergraduate psychology students (beginners) and in graduated psychologists (advanced learners). During fMRI scanning, participants were presented with words denoting scientific psychological concepts within a lexical decision task (e.g. "conditioning", "habituation"). The individual semantic property content of each concept was related to brain activity during abstract concept processing. In both beginners and advanced learners, visual and motor properties activated brain regions also involved in perception and action, while mental state properties increased activity in brain regions also recruited by emotional-social scene observation. Only in advanced learners, social constellation properties elicited brain activity overlapping with emotional-social scene observation. In line with grounded cognition approaches, the present results highlight the importance of experiential information for constituting the meaning of abstract scientific concepts during the course of academic training.

Differential temporo-spatial pattern of electrical brain activity during the processing of abstract concepts related to mental states and verbal associations.

Refined grounded cognition accounts propose that abstract concepts might be grounded in brain circuits involved in mentalizing. In the present event-related potential (ERP) study, we compared the time course of neural processing in response to semantically predefined abstract mental states and verbal association concepts during a lexical decision task. In addition to scalp ERPs, source estimates of underlying volume brain activity were determined to reveal spatio-temporal clusters of greater electrical brain activity to abstract mental state vs. verbal association concepts, and vice versa. Source estimates suggested early (onset 194 ms), but short-lived enhanced activity (offset 210 ms) to verbal association concepts in left occipital regions. Increased occipital activity might reflect retrieval of visual word form or access to visual conceptual features of associated words. Increased estimated source activity to mental state concepts was obtained in visuo-motor (superior parietal, pre- and postcentral areas) and mentalizing networks (lateral and medial prefrontal areas, insula, precuneus, temporo-parietal junction) with an onset of 212 ms, which extended to later time windows. The time course data indicated two processing phases: An initial conceptual access phase, in which linguistic and modal brain circuits rapidly process features depending on their relevance, and a later conceptual elaboration phase, in which elaborative processing within feature-specific networks further refines the concept. This study confirms the proposal that abstract concepts are based on representations in distinct neural circuits depending on their semantic feature content. The present research also highlights the importance of investigating sets of abstract concepts with a defined semantic content.

Time course of brain activity during the processing of motor- and vision-related abstract concepts: flexibility and task dependency.

Grounded cognition theories assume that conceptual processing depends on modality-specific brain systems in a context-dependent fashion. Although the relation of abstract concepts to modality-specific systems is less obvious than for concrete concepts, recent behavioral and neuroimaging studies indicated a foundation of abstract concepts in vision and action. However, due to their poor temporal resolution, neuroimaging studies cannot determine whether sensorimotor activity reflects rapid access to conceptual information or later conceptual processes. The present study therefore assessed the time course of abstract concept processing using event-related potentials (ERPs) and compared ERP responses to abstract concepts with a strong relation to vision or action. We tested whether possible ERP effects to abstract word categories would emerge in early or in later time windows and whether these effects would depend on the depth of the conceptual task. In Experiment 1, a shallow lexical decision task, early feature-specific effects starting at 178 ms were revealed, but later effects beyond 300 ms were also observed. In Experiment 2, a deep conceptual decision task, feature-specific effects with an onset of 22 ms were obtained, but effects again extended beyond 300 ms. In congruency with earlier neuroimaging work, the present feature-specific ERP effects suggest a grounding of abstract concepts in modal brain systems. The presence of early and late feature-specific effects indicates that sensorimotor activity observed in neuroimaging experiments may reflect both rapid conceptual and later post-conceptual processing. Results furthermore suggest that a deep conceptual task accelerates access to conceptual sensorimotor features, thereby demonstrating conceptual flexibility.

The grounding of abstract concepts in the motor and visual system: An fMRI study.

The grounding of concepts in the sensorimotor brain systems is controversially discussed. Grounded cognition models propose that concepts are represented in modality-specific sensorimotor, but also emotional and introspective brain areas depending on specific experiences during concept acquisition. Accumulating evidence suggests that concrete concepts are closely linked to modality-specific systems, whereas the mere existence of abstract concepts seems to contradict grounded cognition approaches. Here, using functional magnetic resonance imaging, we adopted a theory-driven approach frequently used for investigating concrete concepts to the domain of abstract concepts: We compared brain activation to abstract concepts with a known motor versus visual feature content as determined by a previous property listing study. Carefully matched motor (e.g., fitness) and visual (e.g., beauty) abstract words were presented to 24 participants along with pseudowords while performing a lexical decision task. Furthermore, participants performed two localizer tasks by actually moving their hands (motor localizer) and by looking at real pictures (visual localizer). Processing of motor abstract words specifically activated frontal and parietal motor areas, whereas processing of visual abstract words specifically elicited higher activity in temporo-occipital visual areas, albeit at a more lenient statistical threshold. According to inclusive masking analyses, this differential activity pattern to motor and visual abstract concepts overlapped with brain activations observed during hand movements (pre- and postcentral gyrus) and object perception (fusiform and lingual gyrus). Thus, consistent with the grounded cognition framework, our results suggest that, similar to concrete concepts, abstract concepts related to action and vision are grounded in modality-specific brain systems typically engaged in actual perception and action depending on their conceptual feature content.

Processing of Action and Sound Verbs in Context: An FMRI Study.

Recent theories propose a flexible recruitment of sensory and motor brain regions during conceptual processing depending on context and task. The present functional magnetic resonance imaging study investigated the influence of context and task on conceptual processing of action and sound verbs. Participants first performed an explicit semantic context decision task, in which action and sound verbs were presented together with a context noun. The same verbs were repeatedly presented in a subsequent implicit lexical decision task together with new action and sound verbs. Thereafter, motor and acoustic localizer tasks were administered to identify brain regions involved in perception and action. During the explicit task, we found differential activations to action and sound verbs near corresponding sensorimotor brain regions. During the implicit lexical decision task, differences between action and sound verbs were absent. However, feature-specific repetition effects were observed near corresponding sensorimotor brain regions. The present results suggest flexible conceptual representations depending on context and task. Feature-specific effects were observed only near, but not within corresponding sensorimotor brain regions, as defined by the localizer tasks. Our results therefore only provide limited evidence in favor of grounded cognition theories assuming a close link between the conceptual and the sensorimotor systems.

Attentional modulation of masked semantic priming by visible and masked task cues.

In contrast to classical theories of cognitive control, recent evidence suggests that cognitive control and unconscious automatic processing influence each other. First, masked semantic priming, an index of unconscious automatic processing, depends on attention to semantics induced by a previously executed task. Second, cognitive control operations (e.g., implementation of task sets indicating how to process a particular stimulus) can be activated by masked task cues, presented outside awareness. In this study, we combined both lines of research. We investigated in three experiments whether induction tasks and presentation of visible or masked task cues, which signal subsequent semantic or perceptual tasks but do not require induction task execution, comparably modulate masked semantic priming. In line with previous research, priming was consistently larger following execution of a semantic rather than a perceptual induction task. However, we observed in experiment 1 (masked letter cues) a reversed priming pattern following task cues (larger priming following cues signaling perceptual tasks) compared to induction tasks. Experiment 2 (visible letter cues) and experiment 3 (visible color cues) showed that this reversed priming pattern depended only on apriori associations between task cues and task elements (task set dominance), but neither on awareness nor on the verbal or non-verbal format of the cues. These results indicate that task cues have the power to modulate subsequent masked semantic priming through attentional mechanisms. Task-set dominance conceivably affects the time course of task set activation and inhibition in response to task cues and thus the direction of their modulatory effects on priming.

Brain Activation During Conceptual Processing of Action and Sound Verbs.

Grounded cognition approaches to conceptual representations postulate a close link between conceptual knowledge and the sensorimotor brain systems. The present fMRI study tested, whether a feature-specific representation of concepts, as previously demonstrated for nouns, can also be found for action- and sound-related verbs. Participants were presented with action- and soundrelated verbs along with pseudoverbs while performing a lexical decision task. Sound-related verbs activated auditory areas in the temporal cortex, whereas action-related verbs activated brain regions in the superior frontal gyrus and the cerebellum, albeit only at a more liberal threshold. This differential brain activation during conceptual verb processing partially overlapped with or was adjacent to brain regions activated during the functional localizers probing sound perception or action execution. Activity in brain areas involved in the processing of action information was parametrically modulated by ratings of action relevance. Comparisons of action- and sound-related verbs with pseudoverbs revealed activation for both verb categories in auditory and motor areas. In contrast to proposals of strong grounded cognition approaches, our study did not demonstrate a considerable overlap of activations for action- and sound-related verbs and for the corresponding functional localizer tasks. However, in line with weaker variants of grounded cognition theories, the differential activation pattern for action- and sound-related verbs was near corresponding sensorimotor brain regions depending on conceptual feature relevance. Possibly, action-sound coupling resulted in a mutual activation of the motor and the auditory system for both action- and sound-related verbs, thereby reducing the effect sizes for the differential contrasts.

The Semantic Content of Abstract Concepts: A Property Listing Study of 296 Abstract Words.

The relation of abstract concepts to the modality-specific systems is discussed controversially. According to classical approaches, the semantic content of abstract concepts can only be coded by amodal or verbal-symbolic representations distinct from the sensory and motor systems, because abstract concepts lack a clear physical referent. Grounded cognition theories, in contrast, propose that abstract concepts do not depend only on the verbal system, but also on a variety of modal systems involving perception, action, emotion and internal states. In order to contribute to this debate, we investigated the semantic content of abstract concepts using a property generation task. Participants were asked to generate properties for 296 abstract concepts, which are relevant for constituting their meaning. These properties were categorized by a coding-scheme making a classification into modality-specific and verbal contents possible. Words were additionally rated with regard to concreteness/abstractness and familiarity. To identify possible subgroups of abstract concepts with distinct profiles of generated features, hierarchical cluster analyses were conducted. Participants generated a substantial proportion of introspective, affective, social, sensory and motor-related properties, in addition to verbal associations. Cluster analyses revealed different subcategories of abstract concepts, which can be characterized by the dominance of certain conceptual features. The present results are therefore compatible with grounded cognition theories, which emphasize the importance of linguistic, social, introspective and affective experiential information for the representation of abstract concepts. Our findings also indicate that abstract concepts are highly heterogeneous requiring the investigation of well-specified subcategories of abstract concepts, for instance as revealed by the present cluster analyses. The present study could thus guide future behavioral or imaging work further elucidating the representation of abstract concepts.

Functional reorganization of the conceptual brain system after deafness in early childhood.

The neurodevelopmental consequences of deafness on the functional neuroarchitecture of the conceptual system have not been intensively investigated so far. Using functional magnetic resonance imaging (fMRI), we therefore identified brain areas involved in conceptual processing in deaf and hearing participants. Conceptual processing was probed by a pictorial animacy decision task. Furthermore, brain areas sensitive to observing verbal signs and to observing non-verbal visual hand actions were identified in deaf participants. In hearing participants, brain areas responsive to environmental sounds and the observation of visual hand actions were determined. We found a stronger recruitment of superior and middle temporal cortex in deaf compared to hearing participants during animacy decisions. This region, which forms auditory cortex in hearing people according to the sound listening task, was also activated in deaf participants, when they observed sign language, but not when they observed non-verbal hand actions. These results indicate that conceptual processing in deaf people more strongly depends on language representations compared to hearing people. Furthermore, additionally enhanced activation in visual and motor areas of deaf versus hearing participants during animacy decisions and a more frequent report of visual and motor features in the property listing task suggest that the loss of the auditory channel is partially compensated by an increased importance of visual and motor information for constituting object knowledge. Hence, our results indicate that conceptual processing in deaf compared to hearing people is more strongly based on the language system, complemented by an enhanced contribution of the visuo-motor system.

Feature-Specific Event-Related Potential Effects to Action- and Sound-Related Verbs during Visual Word Recognition.

Grounded cognition theories suggest that conceptual representations essentially depend on modality-specific sensory and motor systems. Feature-specific brain activation across different feature types such as action or audition has been intensively investigated in nouns, while feature-specific conceptual category differences in verbs mainly focused on body part specific effects. The present work aimed at assessing whether feature-specific event-related potential (ERP) differences between action and sound concepts, as previously observed in nouns, can also be found within the word class of verbs. In Experiment 1, participants were visually presented with carefully matched sound and action verbs within a lexical decision task, which provides implicit access to word meaning and minimizes strategic access to semantic word features. Experiment 2 tested whether pre-activating the verb concept in a context phase, in which the verb is presented with a related context noun, modulates subsequent feature-specific action vs. sound verb processing within the lexical decision task. In Experiment 1, ERP analyses revealed a differential ERP polarity pattern for action and sound verbs at parietal and central electrodes similar to previous results in nouns. Pre-activation of the meaning of verbs in the preceding context phase in Experiment 2 resulted in a polarity-reversal of feature-specific ERP effects in the lexical decision task compared with Experiment 1. This parallels analogous earlier findings for primed action and sound related nouns. In line with grounded cognitions theories, our ERP study provides evidence for a differential processing of action and sound verbs similar to earlier observation for concrete nouns. Although the localizational value of ERPs must be viewed with caution, our results indicate that the meaning of verbs is linked to different neural circuits depending on conceptual feature relevance.

Handwriting or Typewriting? The Influence of Pen- or Keyboard-Based Writing Training on Reading and Writing Performance in Preschool Children.

Digital writing devices associated with the use of computers, tablet PCs, or mobile phones are increasingly replacing writing by hand. It is, however, controversially discussed how writing modes influence reading and writing performance in children at the start of literacy. On the one hand, the easiness of typing on digital devices may accelerate reading and writing in young children, who have less developed sensory-motor skills. On the other hand, the meaningful coupling between action and perception during handwriting, which establishes sensory-motor memory traces, could facilitate written language acquisition. In order to decide between these theoretical alternatives, for the present study, we developed an intense training program for preschool children attending the German kindergarten with 16 training sessions. Using closely matched letter learning games, eight letters of the German alphabet were trained either by handwriting with a pen on a sheet of paper or by typing on a computer keyboard. Letter recognition, naming, and writing performance as well as word reading and writing performance were assessed. Results did not indicate a superiority of typing training over handwriting training in any of these tasks. In contrast, handwriting training was superior to typing training in word writing, and, as a tendency, in word reading. The results of our study, therefore, support theories of action-perception coupling assuming a facilitatory influence of sensory-motor representations established during handwriting on reading and writing.

Unconscious automatic brain activation of acoustic and action-related conceptual features during masked repetition priming.

Classical theories of semantic memory assume that concepts are represented in a unitary amodal memory system. In challenging this classical view, pure or hybrid modality-specific theories propose that conceptual representations are grounded in the sensory-motor brain areas, which typically process sensory and action-related information. Although neuroimaging studies provided evidence for a functional-anatomical link between conceptual processing of sensory or action-related features and the sensory-motor brain systems, it has been argued that aspects of such sensory-motor activation may not directly reflect conceptual processing but rather strategic imagery or postconceptual elaboration. In the present ERP study, we investigated masked effects of acoustic and action-related conceptual features to probe unconscious automatic conceptual processing in isolation. Subliminal feature-specific ERP effects at frontocentral electrodes were observed, which differed with regard to polarity, topography, and underlying brain electrical sources in congruency with earlier findings under conscious viewing conditions. These findings suggest that conceptual acoustic and action representations can also be unconsciously accessed, thereby excluding any postconceptual strategic processes. This study therefore further substantiates a grounding of conceptual and semantic processing in action and perception.

Masked priming of conceptual features reveals differential brain activation during unconscious access to conceptual action and sound information.

Previous neuroimaging studies suggested an involvement of sensory-motor brain systems during conceptual processing in support of grounded cognition theories of conceptual memory. However, in these studies with visible stimuli, contributions of strategic imagery or semantic elaboration processes to observed sensory-motor activity cannot be entirely excluded. In the present study, we therefore investigated the electrophysiological correlates of unconscious feature-specific priming of action- and sound-related concepts within a novel feature-priming paradigm to specifically probe automatic processing of conceptual features without the contribution of possibly confounding factors such as orthographic similarity or response congruency. Participants were presented with a masked subliminal prime word and a subsequent visible target word. In the feature-priming conditions primes as well as targets belonged to the same conceptual feature dimension (action or sound, e.g., typewriter or radio) whereas in the two non-priming conditions, either the primes or the targets consisted of matched control words with low feature relevance (e.g., butterfly or candle). Event-related potential analyses revealed unconscious feature-specific priming effects at fronto-central electrodes within 100 to 180 ms after target stimulus onset that differed with regard to topography and underlying neural generators. In congruency with previous findings under visible stimulation conditions, these differential subliminal ERP feature-priming effects demonstrate an unconscious automatic access to action versus sound features of concepts. The present results therefore support grounded cognition theory suggesting that activity in sensory and motor areas during conceptual processing can also occur unconsciously and is not mandatorily accompanied by a vivid conscious experience of the conceptual content such as in imagery.

Losing the sound of concepts: damage to auditory association cortex impairs the processing of sound-related concepts.

Conceptual knowledge is classically supposed to be abstract and represented in an amodal unitary system, distinct from the sensory and motor brain systems. A more recent embodiment view of conceptual knowledge, however, proposes that concepts are grounded in distributed modality-specific brain areas which typically process sensory or action-related object information. Recent neuroimaging evidence suggested the significance of left auditory association cortex encompassing posterior superior and middle temporal gyrus in coding conceptual sound features of everyday objects. However, a causal role of this region in processing conceptual sound information has yet to be established. Here we had the unique chance to investigate a patient, JR, with a focal lesion in left posterior superior and middle temporal gyrus. To test the necessity of this region in conceptual and perceptual processing of sound information we administered four different experimental tasks to JR: Visual word recognition, category fluency, sound recognition and voice classification. Compared with a matched control group, patient JR was consistently impaired in conceptual processing of sound-related everyday objects (e.g., "bell"), while performance for non-sound-related everyday objects (e.g., "armchair"), animals, whether they typically produce sounds (e.g., "frog") or not (e.g., "tortoise"), and musical instruments (e.g., "guitar") was intact. An analogous deficit pattern in JR was also obtained for perceptual recognition of the corresponding sounds. Hence, damage to left auditory association cortex specifically impairs perceptual and conceptual processing of sounds from everyday objects. In support of modality-specific theories, these findings strongly evidence the necessity of auditory association cortex in coding sound-related conceptual information.