Interactions between visual and semantic processing during object recognition revealed by modulatory effects of age of acquisition

The age of acquisition (AoA) of objects and their names is a powerful determinant of processing speed in adulthood, with early-acquired objects being recognized and named faster than late-acquired objects. Previous research using fMRI (Ellis et al., 2006. Traces of vocabulary acquisition in the brain: evidence from covert object naming. NeuroImage 33, 958–968) found that AoA modulated the strength of BOLD responses in both occipital and left anterior temporal cortex during object naming. We used magnetoencephalography (MEG) to explore in more detail the nature of the influence of AoA on activity in those two regions. Covert object naming recruited a network within the left hemisphere that is familiar from previous research, including visual, left occipito-temporal, anterior temporal and inferior frontal regions. Region of interest (ROI) analyses found that occipital cortex generated a rapid evoked response (~ 75–200 ms at 0–40 Hz) that peaked at 95 ms but was not modulated by AoA. That response was followed by a complex of later occipital responses that extended from ~ 300 to 850 ms and were stronger to early- than late-acquired items from ~ 325 to 675 ms at 10–20 Hz in the induced rather than the evoked component. Left anterior temporal cortex showed an evoked response that occurred significantly later than the first occipital response (~ 100–400 ms at 0–10 Hz with a peak at 191 ms) and was stronger to early- than late-acquired items from ~ 100 to 300 ms at 2–12 Hz. A later anterior temporal response from ~ 550 to 1050 ms at 5–20 Hz was not modulated by AoA. The results indicate that the initial analysis of object forms in visual cortex is not influenced by AoA. A fastforward sweep of activation from occipital and left anterior temporal cortex then results in stronger activation of semantic representations for early- than late-acquired objects. Top-down re-activation of occipital cortex by semantic representations is then greater for early than late acquired objects resulting in delayed modulation of the visual response

The lexical nature of alpha-beta oscillations in context-driven word production

In context-driven word production, picture naming is faster following constrained than neutral sentential contexts (e.g., “The farmer milked the… [picture]” vs. “The child drew a… [picture]”, followed by the picture of a cow), suggesting conceptual-lexical pre-activation of the target response. Power decreases in the alpha-beta oscillatory band (8–25 Hz) are consistently found for constrained relative to neutral contexts prior to picture onset, when conceptual and lexical retrieval is ongoing. However, it remains a matter of debate whether the alpha-beta power decreases reflect (low-level) expectations of the visual input, conceptual and lexical retrieval, or motor preparation. The present study aimed at investigating the lexical-semantic nature of alpha-beta oscillations. Participants performed context-driven picture naming with constrained and neutral contexts. In addition, an auditory distractor word was presented before picture onset. Distractors were either semantically related (e.g., “goat”) or unrelated (e.g., “bean”) to the picture to be named. Picture naming was faster with constrained than neutral contexts. Distractor type did not affect naming latencies nor the behavioural context effect. In the oscillatory brain responses, the context-related alpha-beta power decreases were observed throughout the pre-picture interval when distractors were semantically unrelated to the picture, in line with previous findings. However, with semantically related distractors, the context effect was delayed until a period after distractor processing. Thus, alpha-beta power seems to be sensitive to the semantic relationship between the distractor word and the picture to be named. We interpret these results as suggesting that alpha-beta power decreases in context-driven word production reflect lexical-semantic retrieval mechanisms

Electrophysiological signatures of conceptual and lexical retrieval from semantic memory

Retrieval from semantic memory of conceptual and lexical information is essential for producing speech. It is unclear whether there are differences in the neural mechanisms of conceptual and lexical retrieval when spreading activation through semantic memory is initiated by verbal or nonverbal settings. The same twenty participants took part in two EEG experiments. The first experiment examined conceptual and lexical retrieval following nonverbal settings, whereas the second experiment was a replication of previous studies examining conceptual and lexical retrieval following verbal settings. Target pictures were presented after constraining and nonconstraining contexts. In the nonverbal settings, contexts were provided as two priming pictures (e.g., constraining: nest, feather; nonconstraining: anchor, lipstick; target picture: BIRD). In the verbal settings, contexts were provided as sentences (e.g., constraining: “The farmer milked a...”; nonconstraining: “The child drew a...”; target picture: COW). Target pictures were named faster following constraining contexts in both experiments, indicating that conceptual preparation starts before target picture onset in constraining conditions. In the verbal experiment, we replicated the alpha-beta power decreases in constraining relative to nonconstraining conditions before target picture onset. No such power decreases were found in the nonverbal experiment. Power decreases in constraining relative to nonconstraining conditions were significantly different between experiments. Our findings suggest that participants engage in conceptual preparation following verbal and nonverbal settings, albeit differently. The retrieval of a target word, initiated by verbal settings, is associated with alpha-beta power decreases. By contrast, broad conceptual preparation alone, prompted by nonverbal settings, does not seem enough to elicit alpha-beta power decreases. These findings have implications for theories of oscillations and semantic memory

Advances in modeling and characterization of human neuromagnetic oscillations

Intracranial electrophysiological measurements as well as electromagnetic recordings from the scalp have shown that oscillatory activity in the human brain plays an important role in sensory and cognitive processing. Communication between distant brain regions seems to be mediated by oscillatory coherence and synchrony. Our brain is both reactive and reflexive: it reacts to changes in the external environment, but it is also influenced by its past and present internal state. On the one hand, task-related or induced modulations of oscillatory activity provide an important marker for cortical excitability and information processing of the reactive brain. On the other hand, spontaneous oscillatory dynamics subserves information processing of the reflexive brain. In this thesis, methods were developed to model and characterize task-related oscillatory changes, as well as spontaneous oscillatory activity measured using magnetoencephalography (MEG). In Publication I, we developed a predictive model to capture the suppression-rebound reactivity of the ~20 Hz mu rhythm originating in the sensorimotor cortex and applied this model to locate the cortical generators of the rhythm from independent measurements. In Publications II and III, we developed temporal and spatial variants of a data-driven method to characterize spatial, temporal, and spectral aspects of spontaneous MEG oscillations. Analysis of complex-valued Fourier coefficients identified well-known rhythms, such as the parieto-occipital ~10-Hz and the rolandic ~20-Hz rhythms consistently across subjects. In Publication IV, we applied independent component analysis to time-frequency representations of cortical current estimates computed from simulated as well as resting-state and naturalistic stimulation data. Group-level analysis of Fourier envelopes also identified the ~20-Hz bilateral sensorimotor network, a subset of the default-mode network at ~8 and ~15 Hz, and lateralized temporal-lobe sources at ~8 Hz. The methods developed here represent important advances in the modeling and characterization of the brain's oscillatory activity measured using non-invasive electrophysiological methods in healthy humans

The impact of MEG source reconstruction method on source-space connectivity estimation: A comparison between minimum-norm solution and beamforming.

Despite numerous important contributions, the investigation of brain connectivity with magnetoencephalography (MEG) still faces multiple challenges. One critical aspect of source-level connectivity, largely overlooked in the literature, is the putative effect of the choice of the inverse method on the subsequent cortico-cortical coupling analysis. We set out to investigate the impact of three inverse methods on source coherence detection using simulated MEG data. To this end, thousands of randomly located pairs of sources were created. Several parameters were manipulated, including inter- and intra-source correlation strength, source size and spatial configuration. The simulated pairs of sources were then used to generate sensor-level MEG measurements at varying signal-to-noise ratios (SNR). Next, the source level power and coherence maps were calculated using three methods (a) L2-Minimum-Norm Estimate (MNE), (b) Linearly Constrained Minimum Variance (LCMV) beamforming, and (c) Dynamic Imaging of Coherent Sources (DICS) beamforming. The performances of the methods were evaluated using Receiver Operating Characteristic (ROC) curves. The results indicate that beamformers perform better than MNE for coherence reconstructions if the interacting cortical sources consist of point-like sources. On the other hand, MNE provides better connectivity estimation than beamformers, if the interacting sources are simulated as extended cortical patches, where each patch consists of dipoles with identical time series (high intra-patch coherence). However, the performance of the beamformers for interacting patches improves substantially if each patch of active cortex is simulated with only partly coherent time series (partial intra-patch coherence). These results demonstrate that the choice of the inverse method impacts the results of MEG source-space coherence analysis, and that the optimal choice of the inverse solution depends on the spatial and synchronization profile of the interacting cortical sources. The insights revealed here can guide method selection and help improve data interpretation regarding MEG connectivity estimation

Roles of Brain Criticality and Multiscale Oscillations in Temporal Predictions for Sensorimotor Processing

Sensorimotor predictions are essential for adaptive behavior. In natural environments, events that demand sensorimotor predictions unfold across many timescales, and corresponding temporal predictions (either explicit or implicit) should therefore emerge in brain dynamics. Neuronal oscillations are scale-specific processes found in several frequency bands. They underlie periodicity in sensorimotor processing and can represent temporal predictions via their phase dynamics. These processes build upon endogenous neural rhythmicity and adapt in response to exogenous timing demands. While much of the research on periodicity in neural processing has focused on subsecond oscillations, these fast-scale rhythms are in fact paralleled by critical-like, scale-free dynamics and fluctuations of brain activity at various timescales, ranging from seconds to hundreds of seconds. In this review, we put forth a framework positing that critical brain dynamics are essential for the role of neuronal oscillations in timing and that cross-frequency coupling flexibly organizes neuronal processing across multiple frequencies.Peer reviewe

From locomotion to dance and back : exploring rhythmic sensorimotor synchronization

Le rythme est un aspect important du mouvement et de la perception de l’environnement. Lorsque l’on danse, la pulsation musicale induit une activité neurale oscillatoire qui permet au système nerveux d’anticiper les évènements musicaux à venir. Le système moteur peut alors s’y synchroniser. Cette thèse développe de nouvelles techniques d’investigation des rythmes neuraux non strictement périodiques, tels que ceux qui régulent le tempo naturellement variable de la marche ou la perception rythmes musicaux. Elle étudie des réponses neurales reflétant la discordance entre ce que le système nerveux anticipe et ce qu’il perçoit, et qui sont nécessaire pour adapter la synchronisation de mouvements à un environnement variable. Elle montre aussi comment l’activité neurale évoquée par un rythme musical complexe est renforcée par les mouvements qui y sont synchronisés. Enfin, elle s’intéresse à ces rythmes neuraux chez des patients ayant des troubles de la marche ou de la conscience.Rhythms are central in human behaviours spanning from locomotion to music performance. In dance, self-sustaining and dynamically adapting neural oscillations entrain to the regular auditory inputs that is the musical beat. This entrainment leads to anticipation of forthcoming sensory events, which in turn allows synchronization of movements to the perceived environment. This dissertation develops novel technical approaches to investigate neural rhythms that are not strictly periodic, such as naturally tempo-varying locomotion movements and rhythms of music. It studies neural responses reflecting the discordance between what the nervous system anticipates and the actual timing of events, and that are critical for synchronizing movements to a changing environment. It also shows how the neural activity elicited by a musical rhythm is shaped by how we move. Finally, it investigates such neural rhythms in patient with gait or consciousness disorders

Exploring the neural entrainment to musical rhythms and meter : a steady-state evoked potential approach

Thèse de doctorat réalisé en cotutelle avec l'Université catholique de Louvain, Belgique (Faculté de médecine, Institut de Neuroscience)Percevoir et synchroniser ses mouvements à une pulsation régulière en musique est une capacité largement répandue chez l’Homme, et fondamentale aux comportements musicaux. La pulsation et la métrique en musique désignent généralement une organisation temporelle périodique perçue à partir de stimuli acoustiques complexes, et cette organisation perceptuelle implique souvent une mise en mouvement périodique spontanée du corps. Cependant, les mécanismes neuraux sous-tendant cette perception sont à l’heure actuelle encore méconnus. Le présent travail a donc eu pour objectif de développer une nouvelle approche expérimentale, inspirée par l’approche électrophysiologique des potentiels évoqués stationnaires, afin d’explorer les corrélats neuraux à la base de notre perception de la pulsation et de la métrique induite à l’écoute de rythmes musicaux. L’activité neurale évoquée en relation avec la perception d’une pulsation a été enregistrée par électroencéphalographie (EEG) chez des individus sains, dans divers contextes : (1) dans un contexte d’imagerie mentale d’une métrique appliquée de manière endogène sur un stimulus auditif, (2) dans un contexte d’induction spontanée d’une pulsation à l’écoute de patterns rythmiques musicaux, (3) dans un contexte d’interaction multisensorielle, et (4) dans un contexte de synchronisation sensorimotrice. Pris dans leur ensemble, les résultats de ces études corroborent l’hypothèse selon laquelle la perception de la pulsation en musique est sous-tendue par des processus de synchronisation et de résonance de l’activité neurale dans le cerveau humain. De plus, ces résultats suggèrent que l’approche développée dans le présent travail pourrait apporter un éclairage significatif pour comprendre les mécanismes neuraux de la perception de la pulsation et des rythmes musicaux, et, dans une perspective plus générale, pour explorer les mécanismes de synchronisation neurale.The ability to perceive a regular beat in music and synchronize to it is a widespread human skill. Fundamental to musical behavior, beat and meter refer to the perception of periodicities while listening to musical rhythms, and usually involve spontaneous entrainment to move on these periodicities. However, the neural mechanisms underlying entrainment to beat and meter in Humans remain unclear. The present work tests a novel experimental approach, inspired by the steady-state evoked potential method, to explore the neural dynamics supporting the perception of rhythmic inputs. Using human electroencephalography (EEG), neural responses to beat and meter were recorded in various contexts: (1) mental imagery of meter, (2) spontaneous induction of a beat from rhythmic patterns, (3) multisensory integration, and (4) sensorimotor synchronization. Our results support the view that entrainment and resonance phenomena subtend the processing of musical rhythms in the human brain. Furthermore, our results suggest that this novel approach could help investigating the link between the phenomenology of musical beat and meter and neurophysiological evidence of a bias towards periodicities arising under certain circumstances in the nervous system. Hence, entrainment to music provides an original framework to explore general entrainment phenomena occurring at various levels, from the inter-neural to the inter-individual level

Anesthetic-induced unresponsiveness: Electroencephalographic correlates and subjective experiences

Anesthetic drugs can induce reversible alterations in responsiveness, connectedness and consciousness. The measures based on electroencephalogram (EEG) have marked potential for monitoring the anesthetized state because of their relatively easy use in the operating room. In this study, 79 healthy young men participated in an awake experiment, and 47 participants continued to an anesthesia experiment where they received either dexmedetomidine or propofol as target-controlled infusion with stepwise increments until the loss of responsiveness. The participants were roused during the constant drug infusion and interviewed. The drug dose was increased to 1.5-fold to achieve a deeper unresponsive state. After regaining responsiveness, the participants were interviewed. EEG was measured throughout the experiment and the N400 event-related potential component and functional and directed connectivity were studied. Prefrontal-frontal connectivity in the alpha frequency band discriminated the states that differed with respect to responsiveness or drug concentration. The net direction of connectivity was frontal-to-prefrontal during unresponsiveness and reversed back to prefrontal-to-frontal upon return of responsiveness. The understanding of the meaning of spoken language, as measured with the N400 effect, was lost along with responsiveness but, in the dexmedetomidine group, the N400 component was preserved suggesting partial preservation of the processing of words during anesthetic-induced unresponsiveness. However, the N400 effect could not be detected in all the awake participants and the choice of analysis method had marked impact on its detection rate at the individual-level. Subjective experiences were common during unresponsiveness induced by dexmedetomidine and propofol but the experiences most often suggested disconnectedness from the environment. In conclusion, the doses of dexmedetomidine or propofol minimally sufficient to induce unresponsiveness do not render the participants unconscious and dexmedetomidine does not completely abolish the processing of semantic stimuli. The local anterior EEG connectivity in the alpha frequency band may have potential in monitoring the depth of dexmedetomidine- and propofol-induced anesthesia.Anesteettien aiheuttama vastauskyvyttömyys: aivosähkökäyräpohjaiset korrelaatit ja subjektiiviset kokemukset Anestesialääkkeillä voidaan saada aikaan palautuvia muutoksia vastauskykyisyydessä, kytkeytyneisyydessä ja tajunnassa. Aivosähkökäyrään (EEG) pohjautuvat menetelmät tarjoavat lupaavia mahdollisuuksia mitata anestesian vaikutusta aivoissa, sillä niitä on suhteellisen helppo käyttää leikkaussalissa. Tässä tutkimuksessa 79 tervettä nuorta miestä osallistui valvekokeeseen ja 47 heistä jatkoi anestesiakokeeseen. Anestesiakokeessa koehenkilöille annettiin joko deksmedetomidiinia tai propofolia tavoiteohjattuna infuusiona nousevia annosportaita käyttäen, kunnes he menettivät vastauskykynsä. Koehenkilöt herätettiin tasaisen lääkeinfuusion aikana ja haastateltiin. Koko kokeen ajan mitattiin EEG:tä, josta tutkittiin N400-herätevastetta sekä toiminnallista ja suunnattua konnektiivisuutta. Prefrontaali-frontaalivälillä mitattu konnektiivisuus alfa-taajuuskaistassa erotteli toisistaan tilat, jotka erosivat vastauskykyisyyden tai lääkepitoisuuden suhteen. Konnektiivisuuden vallitseva suunta oli frontaalialueilta prefrontaalialueille vastauskyvyttömyyden aikana, mutta se kääntyi takaisin prefrontaalisesta frontaaliseen kulkevaksi koehenkilöiden vastauskyvyn palatessa. N400-efektillä mitattu puhutun kielen ymmärtäminen katosi vastauskyvyn menettämisen myötä. Deksmedetomidiiniryhmässä N400-komponentti säilyi, mikä viittaa siihen, että anesteettien aiheuttaman vastauskyvyttömyyden aikana sanojen prosessointi voi säilyä osittain. Yksilötasolla N400-efektiä ei kuitenkaan havaittu edes kaikilla hereillä olevilla henkilöillä, ja analyysimenetelmän valinnalla oli suuri vaikutus herätevasteen havaitsemiseen. Subjektiiviset kokemukset olivat yleisiä deksmedetomidiinin ja propofolin aiheuttaman vastauskyvyttömyyden aikana, mutta kokemukset olivat usein ympäristöstä irtikytkeytyneitä. Yhteenvetona voidaan todeta, että deksmedetomidiini- ja propofoliannokset, jotka juuri ja juuri riittävät aikaansaamaan vastauskyvyttömyyden, eivät aiheuta tajuttomuutta. Deksmedetomidiini ei myöskään täysin estä merkityssisällöllisten ärsykkeiden käsittelyä. Frontaalialueen sisällä EEG:llä mitattu konnektiivisuus alfataajuuskaistassa saattaa olla tulevaisuudessa hyödyllinen menetelmä deksmedetomidiini- ja propofolianestesian syvyyden mittaamiseksi