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Madden-Lombardi, C.

Publications and source records attributed to Madden-Lombardi, C..

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Neural activation down to the spinal cord during action language? A transcranial magnetic stimulation and peripheral nerve stimulation study.

Language comprehension is increasingly recognized as extending beyond the traditional linguistic system to engage motor and perceptual processes. This perspective is supported by numerous studies demonstrating that understanding action-related words often induces behavioral and neurophysiological changes in the motor system. However, it remains unclear whether the influence of action language on the motor system is restricted to cortical regions, or whether it also extends to spinal structures, as observed during motor imagery. To address this, we used transcranial magnetic stimulation and peripheral nerve stimulation to assess corticospinal excitability and cortico-motoneuronal transmission, respectively. Fifteen healthy and right-handed volunteers participated in four conditions: (i) rest, (ii) kinesthetic motor imagery of finger and wrist flexion, (iii) reading action sentences, and (iv) reading non-action sentences. As anticipated, corticospinal excitability increased during both kinesthetic motor imagery and action reading compared to rest. Interestingly, while kinesthetic motor imagery also led to the expected increase in cortico-motoneuronal transmission, no such modulation occurred during action reading. These findings suggest that action reading do not modulate the excitability of high-threshold motoneurons at the spinal level, contrary to motor imagery. Further investigation is needed to test whether action reading activate lower-threshold spinal structures, such as interneurons involved in spinal pre-synaptic inhibition.

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Mental simulations and action language are impaired in individuals with aphantasia

Action reading is thought to engage motor simulations, yielding modulations in activity of motor-related cortical regions, and contributing to action language comprehension. To test these ideas, we measured 1) corticospinal excitability during action reading, and 2) reading comprehension ability, in individuals with normal and impaired imagery (i.e., phantasia and aphantasia, respectively). Thirty-four participants (17 phantasic and 17 aphantasic) were asked to read manual action sentences. By means of transcranial magnetic stimulation, we triggered motor-evoked potentials (MEP) in the target right index finger. MEP amplitude, a marker of corticospinal excitability, increased during action reading relative to rest for phantasic individuals, but not for aphantasic individuals. This result provides neurophysiological evidence that individuals living with aphantasia present a real neurophysiological deficit in motor system engagement during action reading. Furthermore, deep-level reading comprehension ability was impaired in individuals with aphantasia, who had difficulty selecting words that best fit the context of sentences. Altogether, these findings support the idea that motor simulations, along with the activation within the motor system, contribute to action language comprehension.

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Explicit and implicit motor simulations are impaired in individuals with aphantasia

Individuals with aphantasia report having difficulties or an inability to generate visual images of objects or events. So far, there is no evidence showing that this condition also impacts the motor system and the generation of motor simulations. We probed the neurophysiological marker of aphantasia during explicit and implicit forms of motor simulation, i.e., motor imagery and action observation, respectively. We tested a group of individuals without any reported imagery deficits (phantasics) as well as a group of individuals self-reporting the inability to form mental images (aphantasics). We instructed the participants to explicitly imagine a maximal pinch movement in the visual and kinesthetic modalities and to observe a video showing a pinch movement. By means of transcranial magnetic stimulation, we triggered motor-evoked potentials (MEP) in the target right index finger. As expected, MEP amplitude, a marker of corticospinal excitability, increased for phantasics during kinaesthetic motor imagery and action observation relative to rest but not during visual motor imagery. Interestingly, MEP amplitude did not increase in either form of motor simulation for the group of aphantasics. This result provides neurophysiological evidence that individuals living with aphantasia have a real deficit in activating the motor system during motor simulations.

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Reading negative action verbs: one or two-step processing within the primary motor cortex?

Controversy persists regarding the representation of negated actions, specifically concerning activation and inhibitory mechanisms in the motor system, and whether this occurs in one or two steps. We conducted two experiments probing corticospinal excitability (CSE) and short-interval intracortical inhibition (SICI) in the primary motor cortex at different latencies while reading affirmative and negative action sentences. Twenty-six participants read action and non-action sentences in affirmative or negative forms. Using transcranial magnetic stimulation, we probed CSE in hand muscles at rest and at several latencies after verb presentation. We observed a greater CSE for action sentences compared to non-action sentences, regardless of verb form. In experiment two, nineteen participants read affirmative and negative action sentences. We measured CSE and SICI at short and long latencies after verb presentation. CSE was greater for affirmative and negative action sentences at both latencies compared to rest. SICI did not change at the short latency but increased at longer latencies, regardless of verb form. Our results lend partial support for a two-step model, as negated actions showed the same motor excitability as affirmed actions with no additional inhibition at early latencies. Later neural differences between affirmative and negative actions may occur outside the primary motor cortex. Significant statementIn two TMS experiments, we probed corticospinal excitability and short-interval intracortical inhibition in the primary motor cortex at different latencies while subjects read affirmative and negative action sentences. Consistent with an embodied view of language comprehension, our results demonstrate that reading about actions indeed activates the motor system, and this for both negative and affirmative sentences. Our results lend partial support for a two-step model of negation, as negated actions showed the same increase in motor excitability as affirmed actions, with no additional inhibition at early latencies. This suggests that the motor system contributes to comprehension by simulating the negated or affirmed action. Later neural differences between affirmative and negative actions may occur outside the primary motor cortex.

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Imagining and reading actions: towards similar motor representations

While action language and motor imagery both engage the motor system, determining whether these two processes indeed share the same motor representations would contribute to better understanding their underlying mechanisms. We conducted two experiments probing the mutual influence of these two processes. In Exp.1, hand-action verbs were presented subliminally, and participants (n=36) selected the verb they thought they perceived from two alternatives. When congruent actions were imagined prior to this task, accuracy significantly increased, i.e. participants were better able to "see" the subliminal verbs. In Exp.2, participants (n=19) imagined hand flexion or extension, while corticospinal excitability was measured via transcranial magnetic stimulation. Corticospinal excitability was modulated by action verbs subliminally presented prior to imagery. Specifically, the typical increase observed during imagery was suppressed after presentation of incongruent action verbs. This mutual influence of action language and motor imagery, both at behavioral and neurophysiological levels, suggests overlapping motor representations.

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The motor cortex wants the full story: The influence of sentence context on corticospinal excitability in action language processing

The reading of action verbs has been shown to activate motor areas, whereby sentence context may serve to either globally strengthen this activation or to selectively sharpen it. To investigate this issue, we manipulated the presence of manual actions and sentence context, assessing the level of corticospinal excitability by means of transcranial magnetic stimulation. We hypothesized that context would serve to sharpen the neural representation of the described actions in the motor cortex, reflected in context-specific modulation of corticospinal excitability. Participants silently read manual action verbs and non-manual verbs, preceded by a full sentence (rich context) or not (minimal context). Transcranial magnetic stimulation pulses were delivered at rest or shortly after verb presentation. The coil was positioned over the cortical representation of the right first dorsal interosseous (pointer finger). We observed a general increase of corticospinal excitability while reading both manual action and non-manual verbs in minimal context, whereas the modulation was action-specific in rich context: corticospinal excitability increased while reading manual verbs, but did not differ from baseline for non-manual verbs. These findings suggest that sentence context sharpens motor representations, activating the motor cortex when relevant and eliminating any residual motor activation when no action is present.

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Beyond the word and image: III. Neurodynamic properties of the semantic network

Understanding the neural process underlying the comprehension of visual images and sentences remains a major open challenge in cognitive neuroscience. We previously demonstrated with fMRI and DTI that comprehension of visual images and sentences describing human activities recruits a common semantic system. The current research tests the hypothesis that this common semantic system will display similar neural dynamics during processing in these two modalities. To investigate these neural dynamics we recorded EEG from naive subjects as they saw simple narratives made up of a first visual image depicting a human event, followed by a second that was either a sequentially coherent narrative follow-up, or not, of the first image. In separate blocks of trials the same protocol was presented using sentences. Analysis of the EEG signal revealed common neural dynamics for semantic processing across image and sentence modalities. Late positive ERPs were observed in response to sequential incoherence for sentences and images, consistent with previous studies that examined coherence in these two modalities separately. Analysis of oscillatory power revealed increased gamma-band activity for sequential coherence, again consistent with previous studies showing gamma increases for coherence and matching in sentence and image processing. Multivariate analysis demonstrated that training a classifier on data from one modality (images or sentences) allowed reliable decoding of the sequential coherence of data from trials in the untrained modality, providing further support for a common underlying semantic system for images and sentences. Processing sequential coherence of successive stimuli is associated with neural dynamics that are common to sentence and visual image modalities and that can be decoded across modalities. These results are discussed in the context of EEG signatures of narrative processing and meaning, and more general neural mechanisms for structure processing.

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