Postdoctoral Projects

En cours

2022 - 2024

Phonology and grain size as organizing principles of the reading brain across scales

Florence Bouhali
Supervisor: Johannes Ziegler , Laboratoire de Psychologie Cognitive , Chotiga Pattamadilok, Laboratoire Parole et Langage

Neural investigations of reading have mainly focused on describing an average generic left-lateralized reading network, with two  dissociated routes for mapping orthography to phonology, and orthography to lexical forms and meaning, in line with behavioral and computational dual route models of reading. To date, variability in the neural substrates of reading has been associated with some features of the writing system, and with readers’ experience and abilities. For instance, reading through grapheme-to-phoneme mapping, i.e. at a fine grain of orthographic processing, results in consistently more left-lateralized activations in regions of the ventral occipito-temporal cortex (vOT) associated with orthographic processing, compared to mapping whole words (or logograms) to phonology. It remains unclear, however, how to reconcile these various neural findings, implicating different neural features, across ages, abilities, languages, and contexts. Here, we propose a unifying view of the reading network, according to which phonological decoding and orthographic grain size are general organizing principles of the reading brain network across multiple scales (cross-linguistic, inter-individual and intra-individual), and across ages. We predict that increased reliance on fine- grained orthographic processing and on phonological decoding is associated with three neural markers: stronger left-lateralization of vOT reading activations, stronger medial than lateral activations in the left vOT (around the putative grapheme area), and increased reliance on phonological regions of the dorsal reading route. To support to this view, we propose to analyze how these three neural markers are associated with phonology and grain size across ages and scales (across languages, across individuals and across stimuli and contexts) in two datasets: an existing longitudinal and multilingual child dataset (SF immersion dataset), and an adult multimodal dataset to be acquired at the ILCB (ILCB adult dataset). This new multimodal dataset will include 1) behavioral measures of phonological demands and reading grain-size, 2) functional MRI investigations of underlying neural correlates (the three markers) and 3) tests of the causal role of the right-vOT in coarse-grained orthographic processing using TMS. With this project, we hope to advance the understanding of the neural underpinnings of reading, by establishing the role of grain size and reliance on phonology as general organizing principles of the reading brain.


En cours

01/10/2022 - 30/09/2024

Tracing the early developmental milestones of empathy though gestures in preverbal infants

Shreejata Gupta
Supervisor: Isabelle Dautriche , Laboratoire de Psychologie Cognitive , Clément François, Laboratoire Parole et Langage

Empathy, the capacity to share emotional experience of others is fundamental to prosocial behaviours such as cooperation and communication in humans. Investigations into the development of empathy in preverbal infants have solely focussed on their responses to other's facial and vocal markers of emotions. These studies show that infants discriminate emotions, possibly based on their external expressions, but remain inconclusive about infants' comprehension of the meaning of emotions and the underlying mental states. In addition, there is little to no work on whether and how preverbal infants may communicate their empathic motivation as a response to other's emotions. In this project, using behavioural and neurophysiological methods, I propose to investigate gestures in preverbal infants as a way to fill  the existing gaps in our knowledge on early developmental milestones of empathy. Gestures are nonverbal communicative signals produced intentionally to convey a conceptual meaning or achieve a social goal. Thus, because they are inherently a medium of communication of concepts, gestures provide an avenue to study at which age infants understand the conceptual meanings of emotions. Since infants are capable of intentional communicating through gestures from 8 months of age, they may begin empathic responding to others emotional states by producing gestures even before speech. The present project proposes thus to be the first systematic investigation of comprehension and production of gestures during emotion-eliciting events. The results generated will contribute to our understanding of the early cognitive development of empathy; additionally, identifying gesture markers of empathy could be applied into further investigations of empathy development in nonhuman primates, using a comparative framework.

En cours

01/09/2022 - 31/08/2024

Investigation of rhythmic skills and rhythmic background benefits on handwriting in children with dyslexia and/or developmental coordination disorder

Margaux Lê
Supervisor: Jover Marianne, PSYClé , Frey Aline, Laboratoire de Neurosciences Cognitives, Danna Jéremy, Laboratoire de Neurosciences Cognitives.

Dysgraphia, that is characterized by an alteration of handwriting, is often found in developmental coordination disorder (DCD) and/or developmental dyslexia (DD) (Afonso et al., 2019; Biotteau et al., 2019). Interestingly, difficulties to perceive and produce rhythm have also been reported in the latter two disorders (Lense et al., 2021) and could contribute to dysgraphia. However, since graphomotor alteration may be different in DCD and DD (Afonso et al., 2019; Jover & Huau, 2021), the relationship between dysgraphia and rhythm could depend on the associated disorder. This investigation has important implications on both diagnosis and remediation perspectives to help children with dysgraphia. Indeed, a growing number of studies have shown that rhythmic interventions could help patients suffering of motor disorders (e.g. Dalla Bella et al., 2017) and language disorders (e.g. Thomson et al., 2013). Clearly, this project aims to study how rhythm is impacted in children with dysgraphia, in specifying their co-morbidity, and then to evaluate the effects of rhythmic background listening on handwriting in these same children. First, rhythm impairments in children with dysgraphia will be characterized using behavioral and EEG measures of rhythm perception and production. It is expected that children with dysgraphia will perform worse than children without dysgraphia on these rhythm tasks. EEG measures would permit to specify the impacted processes, based on measuring beat-related steady-state evoked potentials (SS-EPs, Nozaradan et al., 2015) in an attempt to disentangle neural sensory and motor contributions to behavioral beat synchronization in children (e.g., Colling et al., 2017). We expect the children to present distinct profiles depending on the presence of DCD and DD, singular or associated. Besides, we expect the handwriting difficulties to coincide with the rhythm ones. The second aim of the project is to investigate the potential benefits of a rhythmic background on handwriting performance in children with dysgraphia. To achieve this second objective, a first study will be conducted in typically developing children (TD) to investigate how rhythmic background presented at different tempi influences handwriting speed and fluency. Then, the benefits of a rhythmic background on handwriting fluency will be formally tested in children with and without dysgraphia. To this aim, children will realize pre-writing and word writing tasks while listening to a rhythmic background with different tempi (matching with the individual handwriting tempo or with an increased/decreased tempo) and in a non-rhythmic background condition. We hypothesize that writing frequency and fluency will improve in the rhythmic background condition. In addition, differential benefits are expected depending on the motor and/or reading disorder associate to dysgraphia.

En cours

01/02/2021 - 31/01/2023

The neural correlates of embodied L2 learning

Ana Zappa
Supervisor: Cheryl Frenck-Mestre, Laboratoire Parole et Langage & Daniel Mestre, Centre de Réalité Virtuelle de la Méditerranée

Within the framework of embodied semantics, the overlap between sensorimotor and language processes could have important implications for second language (L2) acquisition. Neurolinguistic studies have established motor-language interactions and behavioral studies have shown that coupling word encoding with corresponding, relevant gestures enhances language learning. The present study will explore these aspects of language processing and acquisition, using virtual reality (VR), to investigate how performing naturalistic actions during the learning of L2 action verbs may enhance mapping between word form and meaning. We will examine whether “embodied learning”, or learning that occurs using specific physical movements that are coherent with the meaning of new words, creates linguistic representations that produce greater motor resonance (activity in the motor cortex), due possibly to stronger and more specific motor traces, compared to a control condition. In addition, we will investigate whether embodied learning leads to better retention; indeed the implication of both linguistic and motor areas should lead to a more complete semantic representation and increased learning. Training will take place over two days using auditory verbs and a VR oculus headset and a hand controller. Native French speakers will learn the same set of L2 verbs in Serbian, in either the “embodied learning” or the control condition. The “embodied learning” group will learn the action verbs using different specific movements to manipulate objects for each verb; the control group will simply point to the virtual objects. Both pre and post training, time-frequency analysis will be carried out on the EEG data to measure mu and beta suppression, associated with motor activation, while participants passively listen to the new verbs. EEG and behavioral accuracy will also be used to assess learning in an audio-visual match-mismatch task. We expect that the “embodied learning” group will show greater motor activation during verb processing post-training, and that this will correlate with improved learning as indexed by a greater N400 effect and improved behavioral results compared to the control condition. If so, this would suggest that embodied learning adds a motor trace to lexical items, which would support theories of embodied semantics.

En cours

01/04/2021 - 31/03/2023

An investigation into the inhibitory mechanisms underlying inner verbal actions

Ladislas Nalborczyk
Supervisor: F.-Xavier Alarios, Laboratoire de Psychologie Cognitive & Marieke Longcamp, Laboratoire de Neurosciences Cognitives.

The main goal of this project is to tackle the problem of motor inhibition during covert speech and imagined typing, where covert speech is considered as the mental imagery of overt speech. Put simply, how can we imagine raising our arm without actually raising our arm? How can we imagine a conversation without actually producing it overtly? What are the cognitive and neural mechanisms that operate in order to prevent motor execution? How (where and when) are these mechanisms neurally implemented? Can we enhance or degrade these inhibitory mechanisms online? These questions and the problem of motor inhibition emerge from the use of concepts such as simulation or emulation to explain the phenomenon of motor imagery. These views suggest that motor imagery, defined as the mental representation of an action, without overt execution, would result from the simulation or emulation of actual execution. However, this raises the question of how it is possible for imagination of action to not lead to actual execution. We will tackle these questions using novel behavioural paradigms and transcranial magnetic stimulation in a series of five experiments.

En cours

01/01/2021 - 31/12/2022

Functional role of oscillatory dynamics in motor cortex during speech perception

Noémie te Rietmolen
Supervisor: Kristof Strijkers, Laboratoire Parole et Langage & Benjamin Morillon, Institut de Neurosciences des Systèmes.

While our knowledge on the brain structures underpinning speech perception has greatly advanced in the last decades, the neurophysiological mechanisms that can explain how humans process speech are still largely unknown.
In particular, influential theories about speech perception do not agree on the role of the motor system (Skipper et al., 2017): Dual-stream theories suggest that the motor system is not crucial (Hickok & Poeppel, 2007; Hickok, 2014), whereas opposing theories ascribe a fundamental role to the motor system in speech perception (Barnaud et al., 2018; Pulvermüller & Fadiga, 2010). A fruitful approach to understand the neural mechanisms underlying speech perception is to investigate cortical oscillations. Cortical oscillations refer to synchronized rhythmic brain activity, which is hypothesized to be important for structuring, binding and consolidating complex information in the cerebral cortex (e.g. Buzsáki & Draguhn, 2004). Given the intriguing possibility that cortical oscillations may offer a link between brain and behavior, in particular for higher-order cognitive processes such as language perception (e.g. Buzsáki, 2010), the current project sets out to investigate how brain oscillations in the motor cortex impact speech comprehension.
The objectives and hypotheses of this project are guided by current observations and proposals regarding the nature of cortical oscillations for (1) the extraction of speech sounds and (2) the extraction of meaning when perceiving language. With regard to speech sound processing (1), it has been suggested that cortical oscillations “provide the [temporal] infrastructure to parse and decode connected speech” (Giraud & Poeppel, 2012). At the level of the auditory cortex, low-frequency neural oscillations entrain to the (quasi-)rhythmic structure of the speech signal and causally contribute to speech comprehension (Peelle, 2018; Riecke et al., 2018; Zoefel et al., 2018). Moreover, neural entrainment to speech is also observed in regions beyond the auditory cortex, and in particular in the motor cortex, at the phrasal (0.6-1.3 Hz), lexical (1.3-3 Hz), and syllabic rates (3.5-4.5 Hz) (e.g. Keitel et al., 2018; Assaneo & Poeppel, 2018). One hypothesis is that such entrainment in these frequencies reflects temporal prediction derived from the temporal regularities presented in speech (e.g. Morillon & Baillet, 2017). In a similar vein, cortical oscillations related to the lexical meaning of spoken words (2) have been observed over auditory and motor cortices in the high-frequency range (beta and gamma; e.g. Pulvermüller et al., 1996; Canolty et al., 2007).
These large-scale synchronizations between fronto-central and superior temporal brain regions are hypothesized to reflect the binding of sensorimotor experiences into lexical categories (e.g. Strijkers, 2016; Garagnani et al., 2017). However, at present the functional role (if any) of such motor oscillatory activity in speech perception remains debated. In the current project, we set out to investigate the exact nature of oscillatory dynamics in the motor cortex for key components of speech perception (i.e. sound- and meaning-extraction) with two complementary studies
that each containing a behavioral and neurophysiological (magnetoencephalography; MEG) part. The behavioral experiments will assess whether activation of the motor cortex improves speech perception (and if so, under which conditions), and the MEG experiments will assess whether these potential behavioral improvements are indeed driven by frequency-specific cortical oscillations and enhanced functional coupling between motor and auditory cortical regions. In this manner, the results of this project may provide valuable insights for the theoretical development of sensorimotor integration during language processing and even highlight that specific oscillatory patterns (different frequency ranges) drive different processes involved in the perception of speech.

En cours

01/10/2020 - 30/09/2022

Growing and learning with laughter

Chiara Mazzocconi
Supervisor: Abdellah Fourtassi, Laboratoire d'Informatique et Systèmes & Beatrice Priego-Valverde, Laurent Prevot, Maud Champagne-Lavau, Laboratoire Parole et Langage.

Laughter is one of the earliest means that an infant has to convey meaning, practicing turn-taking, attention sharing, directing other's attention and contribute to interaction at the same level of an adult. Through devel- opment its use becomes more and more sophisticated both from a semantic and pragmatic perspective, being closely entangled with language production to convey meaning multimodally. Laughter both when occurring in relation to humour or not, can give us important insights into the child's cognitive, linguistic and pragmatic development on di erent levels of observation. Nevertheless, there is a dearth of research on the development of laughter, especially in interaction. The goal of the current project is to deepen our characterization of laughter development longitudinally in itself and in relation to speech, language, humour and pragmatic abilities. I propose to conduct cross-linguistic corpus studies in typically developing children, and compare the data collected with children with atypical language or pragmatic development to test the hypothesis that laughter can be an
early diagnostic concerning language and pragmatic development even before gesture and speech emerge. In addition to corpus studies, I will run experiments that will contribute to deepening our understanding of laughter perception in development and autism, as well as sheding light on the role of laughter in pragmatic reasoning and irony interpretation. I believe that a deeper understanding of laughter production and processing in clinical populations can give us insights about laughter behaviour in itself and about the clinical condition investigated.


01/10/2019 - 30/09/2021

Multimodal study of functional organization of the Visual Word Form Area and its communication with the spoken language syste

Shuai Wang
Supervisor: Chotiga Pattamadilok, Anne-Sophie Dubarry, Laboratoire Parole et Langage & Dr. Agnès Trébuchon,
Collaboration : Prof. Kennett Pugh, Haskins Lab.

At ILCB, the goal of my research is to investigate 1) the fine-scale spatial organization of functionally segregated neuronal populations within the visual word form area (VWFA) and 2) the activation time course of the VWFA in response to speech as well as the temporal dynamics of the communication between this area and the spoken language network. Broadly, I’m interested in the functional integration and segregation of the language system.

En cours

01/10/2019 - 30/09/2023


Etienne Thoret
Supervisor: Richard Kronland-Martinet, Laboratoire Perception Représentations Image Son Musique & Valentin Emiya, Laboratoire d'Informatique et Systèmes & Stéphane Ayache, Laboratoire d'Informatique et Systèmes
Collaboration : Bruno Torrésani, Institut de Mathématiques de Marseille

I'm a sound and hearing researcher interested to decypher the neurocomputational bases of audition. My researches combine advanced mathematical modeling of sound signals with statistical learning techniques, behavioral testing and neuroinspired techniques in order understand how these processes guide human communication and behaviour.
I'm currently post-doc between the Perception, Representation, Image, Sound, Music lab (PRISM) and the Laboratoire d'Informatique & Systèmes lab (LIS) in Marseille through the Institute of Language Communication & the Brain (ILCB) of Aix-Marseille University. I'm advised by Richard Kronland-Martinet (PRISM) and Valentin Emiya & Stéphane Ayache (LIS).
I'm grateful having been advised by Daniel Pressnitzer & Christian Lorenzi at the Ecole Normale Supérieure de Paris, Stephen McAdams & Philippe Depalle at McGill University in Montreal, and by Sølvi Ystad & Mistuko Aramaki at the CNRS Mechanics and Acoustic Lab in Marseille.

Selected publications :

  • Thoret, E., Andrillon, T., Leger, D., Pressnitzer, D. (2020) Probing machine-learning classifiers using noise, bubbles, and reverse correlation, bioRxiv, bioRxiv 2020.06.22.165688, 10.1101/2020.06.22.165688
  • Thoret, E., Caramiaux, B., Depalle, P., McAdams, S. (In press) Learning metrics on spectrotemporal modulations reveals the perception of musical instrument timbre, Nature Human Behaviour. 10.1038/s41562-020-00987-5
  • Thoret, E., Depalle, P., McAdams, S. (2016) Perceptually salient spectro-temporal modulations for recognition of sustained musical instruments. The Journal of the Acoustical Society of America, 140(6), EL478-EL483. 10.1121/1.4971204
  • Thoret, E., Aramaki, M., Kronland-Martinet, R., Velay, J. L., Ystad, S. (2014) From Sound to Shape: Auditory Perception of Drawing Movements, Journal of Experimental Psychology: Human Perception and Performance, 40(3), 983-994.
  • Thoret, E., Aramaki, M., Bringoux L., Ystad S., Kronland-Martinet R. (2016) Seeing circles and drawing ellipses: when sound biases reproduction of visual motion. PLoS one, 11(4):e0154475. 10.1371/journal.pone.0154475



01/01/2019 - 31/12/2021

Nested cortical organisation models for human and non-human primate inter-species comparisons. Application to a phylogenetic study of the primate cognitive vocal control brain network

Kep Kee Loh
Supervisor: Olivier Coulon, Institut de Neurosciences de la Timone, Adrien Megerditchian, Laboratoire de Psychologie Cognitive

Inter-species comparisons of brain organization between human and non-human primates can provide insights into how uniquely human abilities, such as speech and language, emerged through primate brain evolution. While brain organization can be described in many ways, we focus primarily on cortical folding patterns, or sulci, which are critical landmarks that are strongly tied to the functional and histological features of the brain.
The first goal of my project is to construct the first cortical sulci models that describe the organisation of brain folding patterns (sulci) in four primate species: macaques, baboons, chimpanzees and humans. On the basis of common/homologous sulci, these models allow the registration of individual brains both, within the same species, and across species, for brain comparisons. The second goal of my project is then to apply these models to study how the primate vocal control brain network has changed across the four primate species to understand how speech and language areas emerge in the human brain.


01/04/2019 - 31/03/2021


Nicole Voges
Supervisor: Andrea Broveli, Institut de Neurosciences de la Timone & Demian Battaglia, Laboratoire de Neurosciences Cognitives

Cognitive function arises from the coordinated activity of neural populations distributed over largescale brain networks. However, it is challenging to understand how specific aspects of neural dynamics translate into operations of information processing, and, ultimately, cognitive functions. To address this question, we combine novel approaches from information theory with computational simulations of canonical neural circuits, emulating well-defined cognitive functions. Specifically, we simulate circuits composed of one or multiple brain areas, each modeled as a 1D ring network of simple rate units. Despite its simplicity, such model can give rise to rich neuronal dynamics [1]. These models can be used to reproduce functions such bottom-up transfer of stimuli, working memory and even top-down attentional modulation [2].
We then apply recent tools from the Information Dynamics framework to simulated data. Information Dynamics is a novel theoretical approach that formalize the decomposition of generic information processing into “primitive” operations of active storage, transfer and modification of information [3]. In particular, we analyze simulated recordings from our models, quantifying how its nonlinear dynamics implement specific mix of these different primitive processing operations, varying depending on the emulated cognitive function. For instance, we show that the neuronal subsets maintaining sensory representations in working memory (via reverberant self-sustained activity) can be revealed by high values of the active Information Storage metric. Or, the integration of top-down signals (mediated by nonlinear interactions between active sub-populations) is detected by increased values of information modification.
Our models thus highlight transparently the capacity of information dynamics metrics to characterize which network units participate to cognition-related information processing, and how they do it. Such capability can be exploited for the analysis of actual human MEG datasets.
1. Roxin, A., Brunel, N., Hansel, D. (2005). Physical Review Letters 94(23), 238103
2. Ardid, S., Wang, X., Compte, A. (2007). J Neurosci 27(32), 222
3. Wibral, M., Priesemann, V., Kay, J., Lizier, J., Phillips, W. (2017). Brain and cognition 112, 25


01/10/2018 - 31/09/2020

Understanding brain-behavior relationships with deep multi-view regression

Akrem Sellami
Supervisor: Sylvain Takerkart, Institut de Neurosciences de la Timone & François-Xavier Dupé, Laboratoire d'Informatique et Systèmes & Hachem Kadri, Laboratoire d'Informatique & Systèmes

En cours

01/01/2020 - 30/04/2022

Intergenerational non-verbal communication through interpersonal motor alignment

Birgit Rauchbauer
Supervisor: Marie-hélène Grosbas, Laboratoire de Neurosciences Cognitives

This project investigates the neural underpinnings of non-verbal communication between adults and adolescents in relation to emotional expressions, using functional magnetic resonance imaging (fMRI). For this we are specifically interested in investigating the alignment of actions between adult and adolescents in the context of the emotions expressed. Also, we will specifically investigate the differences of emotion perception expressed by different generations. This project will elucidate the neural mechanisms underlying intergenerational non-verbal communication.
Improvement of motor speech sequence learning through interpersonal motor alignment
Synchronous activity, the temporal matching of body movements and vocal expressions (e.g., singing), may have a positive effect on learning. Yet, it has not been investigated whether this extends to language learning, in particular the learning of motor speech sequences. Also, as we pointed out recently, while research on synchrony is well established in adult, infant and children studies, there is a gap for adolescent populations (Rauchbauer & Grosbras, 2020). This project will thus investigate whether synchronous activity (joint laughter, physical activity or joint singing), online or in a laboratory setting, improves task performance in a motor speech learning task, in adolescents and adults.