Newborns are attracted to voices, faces and social gestures. Paying attention to these social cues in everyday life helps infants and young children learn how to interact with others. During this period of development, a network of connections forms between different parts of the brain that helps children to understand other people’s social behaviors.

During their first year of life, infants who later develop autism spectrum disorders (ASD) pay less attention to social cues. This early indifference to these important signals leads to social deficits in children with ASD. They are less able to understand other people’s behaviors or engage in typical social interactions. It’s not yet clear why children with ASD are less attuned to social cues. But is likely that the development of brain networks essential for understanding social behavior suffers as a result. Studying how such networks develop in typical very young children and those with ASD may help scientist learn more.

Now, Sperdin et al. confirm there are differences in the social brain-networks of very young children with ASD compared with their typical peers. In the experiment, 3-year-old children with ASD and without watched videos of other children playing, while Sperdin et al. recorded what they looked at and what happened in their brains. Eyemovements were measured with a tracker, and the brain activity was recorded using an electroencephalogram (EEG), which uses sensors placed on the scalp to measure electrical signals.

What children with ASD looked at was different than their typical peers, and these differences corresponded with alterations in the brain networks that process social information. Children with ASD who had less severe symptoms had stronger activity in these brain networks. What they looked at also was more similar to typical children. This suggests less severely affected children with ASD may be able to compensate that way.

Identifying ASD-like behaviors and brain differences early in life may help scientists to better understand what causes the condition. It may also help clinicians provide more individualized therapies early in life when the brain is most adaptable. Long-term studies of these brain-network differences in children with ASD are necessary to better understand how therapies can influence these changes.

Early preferential attention to social cues is a fundamental mechanism that facilitates interactions with other human beings. During the third trimester of gestation, the human foetus is already sensitive to both voices (DeCasper and Spence, 1986) and face-like stimuli (Reid et al., 2017). Newborns orient to biological motion (Simion et al., 2008) and prefer their mothers’ voices to those of other females (DeCasper and Fifer, 1980). Infants as young as two weeks imitate faces and human gestures (Meltzoff and Moore, 1977). The orientation to and interaction with social cues during infancy drives the later acquisition of social communication skills during toddler and preschool years. As function of experience, the repeated exposure leads to the progressive emergence of adaptive interactions with conspecifics. Alongside, the brain develops a network of cerebral regions specialized in understanding the social behaviours of others. This network includes the orbitofrontal and medial prefrontal cortices, the superior temporal cortex, the temporal poles, the amygdala, the precuneus, the temporo-parietal junction, the anterior cingulate cortex (ACC) and the insula (Brothers, 1990; Frith and Frith, 2010; Adolphs, 2009; Blakemore, 2008). Collectively, these areas form the social brain and are all implicated to some extent in processing social cues and encoding human social behaviours (Brothers, 1990; Frith and Frith, 2010; Adolphs, 2009; Blakemore, 2008).

Autism is a life-long lasting, highly prevalent neurodevelopmental disorder that affects core areas of cognitive and adaptive function, communication and social interactions (Christensen et al., 2016). A common observation in infants later diagnosed with ASD is the presence of less sensitivity and diminished preferential attention to social cues during the first year of life (Osterling and Dawson, 1994). Toddlers with ASD orient preferentially to non-social contingencies (Klin et al., 2009). Indifference to voices (Sperdin and Schaer, 2016) and faces (Grelotti et al., 2002) in ASD leads to deficits in the development of adapted social interactions with others and to difficulties in understanding human behaviours. It is not established why children with ASD show insensitivity to stimuli with social contingencies at early developmental stages, but this apparent indifference to social cues ultimately hinders the normal development of the social brain network or parts thereof (Pelphrey et al., 2011; Gotts et al., 2012). Some authors propose that deficits in the development of social cognition (such as learning to attribute mental states to others, 'theory of mind' [Frith, 1989]) and/or in sensory processing (Dinstein et al., 2012) prevent children with ASD to actively and appropriately engage with social stimuli. Another hypothesis suggests that they have difficulties building up stimulus-reward contingencies for social stimuli, due to a reduced motivation to attend and engage with them. Regardless of the reasons behind reduced social orienting, diminished interaction and exposure to social stimuli may in turn impede the development of the social brain at early developmental stages in ASD (Chevallier et al., 2012; Dawson et al., 2004).

Evidence remains limited for brain network alterations in response to socially meaningful stimuli in ASD during the period spanning the toddler to preschool years, partly because the acquisition of data during that age period is extremely challenging (Raschle et al., 2012). However, studying very young children with ASD, closer to their diagnosis, is all the more important when recent findings suggest the presence of major developmental changes in large-scale brain networks between adults and younger individuals with ASD (Nomi and Uddin, 2015). Currently, it remains unclear how autism affects the development of the functional brain networks implicated in the processing of socially meaningful information at early developmental stages. A better delineation of the timing and nature of the neurodevelopmental alterations associated with core social deficits in autism may in turn help to improve therapeutic strategies.

Electroencephalography (EEG) is as a powerful non-invasive method to study atypical brain responses to social stimuli in clinical paediatric populations with ASD. For example, surface-based experiments have reported aberrant evoked potentials in response to dynamic eye gaze in infants at high-risk for ASD (Elsabbagh et al., 2012) or to speech stimuli in toddlers with ASD (Kuhl et al., 2013) with differences in resting EEG power in infants at high-risk for ASD (Tierney et al., 2012). Whilst useful, most of the EEG experiments performed on very young children with ASD (younger than four years) have been done with few electrodes only and the analysis restricted to the sensor space. Therefore, hypothetical alterations in the functional brain networks underlying the processing of social stimuli remain to be determined for that age period in ASD.

Here, we recorded high-density EEG and high resolution eye-tracking in toddlers and preschoolers with ASD and their TD peers as they watched naturalistic and ecologically valid dynamic social movies. Using data-driven methods, we first investigated whether the visual exploration behaviour was atypical in toddlers and preschoolers with ASD using kernel density distribution estimations. Then, we explored whether their ongoing source-space directed functional connectivity was altered compared to their TD peers using Granger-causal modelling applied to the EEG source signals. This method estimates brain connectivity in the frequency domain. It identifies which brain regions are the key drivers of information flow in a brain network and directional relationships between brain regions that belong to a network (Coito et al., 2016b). This approach has been applied to study connectivity alterations using intra-cranial recordings (Wilke et al., 2009; van Mierlo et al., 2013; van Mierlo et al., 2011) as well as source imaging based on EEG recordings in clinical populations (Ding et al., 2007; Coito et al., 2016a; Coito et al., 2015; Coito et al., 2016b) and in healthy human participants (Astolfi et al., 2007; Hu et al., 2012; Plomp et al., 2015b). Finally, we looked for relationships between directed functional connectivity measures, visual exploration behaviour and clinical phenotype. As toddlers with ASD have less preferential attention for social cues, we hypothesized that they would show both a different visual exploration behaviour of the dynamic social images and altered directed functional connectivity patterns in brain regions involved in processing social information compared to their TD peers.

Abnormal processing of social cues is a hallmark of ASD (Chevallier et al., 2012; Dawson et al., 2004; Dichter et al., 2009; Elsabbagh et al., 2012; Gotts et al., 2012; Greene et al., 2011; Klin et al., 2009; Pelphrey et al., 2011). However, evidence for alterations of social brain networks at early stages of ASD is scant. Using data-driven methods, we observed aberrant gaze patterns together with frequency specific alterations in the directed functional connectivity in toddlers and preschoolers with ASD when exploring dynamic social stimuli compared to their TD peers. These differences manifested as increased driving and hyper-connectivity in the theta frequency band from nodes that include the right orbital part of the superior frontal gyrus, the bilateral orbital parts of the middle frontal gyri, the right middle cingulate gyrus, the left superior occipital gyrus and the left STG. For the alpha band, we found increased driving from the right orbital part of the middle frontal gyrus and the left cuneus and decreased connectivity from the right STG. To the best of our knowledge, this is the first evidence indicating concomitant alterations in the visual exploration of dynamic social images and in the directed functional connectivity involving key nodes of the social brain (Brothers, 1990; Frith and Frith, 2010; Adolphs, 2009; Blakemore, 2008) at early stages of ASD.

The results indicate that the highest information transfer (i.e. summed outflow) occurs at the global brain level in the theta band (4–7 Hz) followed by the alpha frequency band (8–12 Hz). As such, our data-driven approach reveals high information transfer in physiologically relevant frequency bands given the young age of our participants. These are, namely, prominent brain rhythms during infancy and toddlerhood (Saby and Marshall, 2012; Orekhova et al., 2006). Throughout development, slow waves modulate attentional brain states, encode specific information and ease communication between neuronal populations (Lopes da Silva, 2013). Theta and alpha bands are thought to underlie different cerebral functions, but are closely related (Klimesch, 1999). During infancy and early childhood, modulations in alpha band activity have been associated with the progressive development of visual attentional networks and inhibition of task-irrelevant brain areas (Orekhova et al., 2001; Stroganova et al., 1999), while theta is thought to play a functional role in memory formation, emotional and cognitive functioning (Saby and Marshall, 2012; Orekhova et al., 2006).

In our experiment, the highest information transfer occurred in the theta band. In very young children, theta modulations have been related to the development of the social brain. For example, surface-based EEG experiments in TD infants report enhanced theta power to social versus non-social stimuli at 12 months (Jones et al., 2015). Theta increases during attention to social stimulation in infants and preschool aged children (Orekhova et al., 2006). Hence, social contingencies modulate theta band activity. Similarly, our results show high information transfer in theta when toddlers and preschoolers visually explore dynamic social stimuli.

The development of attentional processes in young children has been associated with modulations in alpha band activity (Orekhova et al., 2001). Conversely, our young participants had to deploy their attentional focus to the dynamic social stimuli. This would explain why high information transfer was also found in the alpha frequency band.

EEG experiments in individuals with ASD show a reduction or an increase in coherence patterns in the theta and/or alpha frequency bands compared to their TD peers at various ages and under different experimental conditions (Schwartz et al., 2017). However, most of the available EEG experiments performed on very young children with ASD and analysis thereof were so far restricted to the scalp surface. As a result, information remains limited regarding the presence of frequency specific alterations within brain regions when young children are exposed to social stimuli. Here, our data-driven source-space approach revealed not only high information transfer in the theta and alpha frequency bands, but also, the involvement of the bilateral medial and the superior orbital frontal regions, the bilateral hippocampi, the bilateral ACC and right amygdala. These areas are implicated in processing social cues and encoding human social behaviours (Brothers, 1990; Frith and Frith, 2010; Adolphs, 2009; Blakemore, 2008).

Our results further indicate the presence of frequency specific alterations in the driving from several brain areas in the toddlers and preschoolers with ASD compared to their TD peers. In the theta band, we found an overall dominant higher driving within several frontal and the cingulate regions. In TD individuals, theta generates within the frontal cortices or the ACC (Asada et al., 1999). In comparison to their TD peers, both these areas develop differently in young toddlers later diagnosed with ASD (Schumann et al., 2010). Accordingly, our results raise the possibility that the brain regions generating theta also follow a different development in the toddlers and preschoolers with ASD. For the alpha band, alteration in the driving was evident from the right orbital part of the middle frontal gyrus. Although experiments during the first three years of life are currently sparse, increased alpha-range EEG connectivity over frontal and central electrodes has recently been reported in high-risk infants who were diagnosed with ASD at 36 month (Orekhova et al., 2014). A magnetoencephalography (MEG) study performed at rest using a source-space approach reported lower coherence in the theta and alpha bands within parietal and occipital regions but their ASD group only included adolescents (Ye et al., 2014). The differences between this specific study and our could stem from either variations in the age groups (adolescents versus toddlers and preschoolers), the stimuli used (grey cross inside a white circle versus dynamic biological visual stimuli) or the methods. More generally, several factors explain discrepancies in brain connectivity results between studies. The type of connectivity measures applied, the approach (surface versus source based), the brain regions analysed and frequency bands examined are variables that influence the results or the age of the participants (Mohammad-Rezazadeh et al., 2016).

Frontal and cingulate areas have been implicated in various complex aspects of social cognition, social reward, social perception and social behaviour (Jonker et al., 2015; Apps et al., 2013). Metabolic changes within the medial prefrontal cortex and the cingulate cortex are correlated with social interaction impairments in childhood ASD (Ohnishi et al., 2000). Several experiments report structural (Patriquin et al., 2016) and functional (Gotts et al., 2012; Patriquin et al., 2016; Greene et al., 2011; Cheng et al., 2015) alterations within these brain areas in school aged children, adolescents and adults with ASD when compared to their TD peers. A recent study described hyper-connectivity within the ACC and bilateral insular cortices in a sample including children aged between seven to 12 years (Uddin et al., 2013a). Some authors propose that the two together form the salience network, whose role is to direct attention to behaviourally-relevant stimuli (Menon and Uddin, 2010). Although we didn’t find differences in the driving from the insula compared to the TD peers, there is an increasing number of evidence showing an abnormal development of the salience network or components thereof in ASD (Uddin, 2015), which may partially explain the limited interest for and engagement with social stimuli that is often observed in individuals with ASD and that constitutes a hallmark of the disorder (Klin et al., 2009; Pelphrey et al., 2011). Accordingly, the toddlers and preschoolers with ASD had a different visual exploration behaviour of the dynamic social stimuli raising the possibility of a reduced interest to visually engage with them. Alternatively, alterations in driving from these regions could partially reflect a reduced motivation to attend and engage with the dynamic social stimuli (Chevallier et al., 2012; Dawson et al., 2004). The alterations in the driving in the alpha band we found here, could also be related to the presence of developmental impairments in attentional networks and/or inhibitory functions (Keehn et al., 2013).

We found higher driving in theta from the left superior temporal and occipital gyri. In the alpha band, we found alterations in the driving from a node in the right STG and the left cuneus in the occipital lobe. Those brain areas are implicated in the processing of biological motion, in analysing the intentions of other people’s actions and self-reflection (Pelphrey and Carter, 2008; Pelphrey et al., 2005; Pelphrey and Morris, 2006; Pelphrey et al., 2004). Our result would suggest that the exploration of the dynamic social visual stimuli that contained biological movements led to altered driving from these brain areas in both frequency bands in the toddlers and preschoolers with ASD compared to their TD peers.

Overall hyper-connectivity seems prevalent in younger populations (that is, infants at high-risk for ASD, toddlers and preschoolers with ASD) while hypo-connectivity is more observed during adolescence and adulthood in ASD (Uddin et al., 2013b). Conversely, a developmental shift occurs in brain growth with an initial period of early brain overgrowth followed by normalization sometime during adolescence (Courchesne et al., 2011). Accordingly, structural white matter connectivity studies also highlight this shift from higher structural connectivity in very young children with ASD to lower connectivity in older children with ASD (Hoppenbrouwers et al., 2014; Conti et al., 2015). Therefore, a global higher-driving and hyper-connectivity from key nodes of the social brain in the theta frequency band in our ASD group is consistent with reports in the literature when considering the very young age of our participants (around 3 years of age on average). For the alpha frequency band, we found alterations in the driving manifesting as both hyper-connectivity from a frontal and an occipital area and under-connectivity from a node in the superior temporal pole. Hence, we found frequency-specific network alterations with distinct patterns of directed functional connectivity in the toddlers and preschoolers with ASD compared to their TD peers. This result is in line with recent experiments indicating the presence of distinct patterns of hyper- and hypo-connectivity between brain regions functionally defined by neural oscillatory activity in children and adolescents with ASD (Ye et al., 2014; Kitzbichler et al., 2015; Datko et al., 2016).

We further explored associations between the driving in the nodes of the network (that is, the summed outflow) and clinical and behavioural phenotypes for each frequency band. We didn’t observe any significant relationships between summed outflow and the ADOS severity scores after FDR correction in either frequency bands. In theta, we observed an increased driving from the median cingulate cortex and the paracentral lobule in the toddlers and preschoolers with ASD who had a more similar visual exploration pattern to their TD peers. Thus, an improved visual exploration pattern of the dynamic social images was related to increased summed outflow in theta from these regions. Moreover, higher summed outflow from the right lingual area was related to better socialization behaviour and leisure and play skills as measured by the VABS-II. Higher summed outflow from the left Heschl's area near the posterior convolutions of the insula and the left rolandic operculum near the circular sulcus of the insula rostrally were positively related to better fine and gross motor skills as measured by the PEP-3. Finally, for the alpha band, we found that higher driving within the left hippocampal area and the left rolandic operculum were positively related to better fine and gross motor skills as measured by the PEP-3. As such, overall increased activity in the theta band within dorsomedial frontal, inferior temporal and insular cortical regions were associated with lower clinical impairment and less atypical gaze patterns whereas increased driving in the alpha band was selectively associated with better motor performance. The presence of hyperactivity within relevant brain region has been interpreted as a possible compensatory mechanism when performing a social target detection task, in adults with ASD at least (Dichter et al., 2009). While to the best of our knowledge, there is currently no other relevant experimental data that addresses this question, we speculate that the overall hyper-driving from these relevant brain regions might be a mechanism to compensate for atypical development of the brain's circuitry over time as higher directed functional connectivity was related improved socialization, motor behaviours and better visual exploration of dynamic social images. However, longitudinal measurements are necessary to fully confirm this interpretation. De facto, the toddlers and preschoolers with ASD who had better gaze performance had better adaptive behaviour, improved global and fine motor skills and enriched interpersonal relationships as measured by the VABS-II and better visual motor imitation skills as measured by the PEP-3. They were also those who had higher summed outflow in several relevant brain areas.

Beyond functional and structural brain alterations reported elsewhere in older children and adults with ASD, our results suggest for the first time, the presence of frequency specific alterations in the driving of information flow from brain areas implicated in social information processing during the viewing of naturalistic dynamic social images in toddlers and preschool with ASD. Furthermore, we show that these frequency specific directed functional connectivity network alterations within regions of the social brain are present at early stages of ASD, justifying further investigation into how early therapeutic interventions targeting social orienting skills may help to remediate social brain development during this critical age period when plasticity is still possible. Longitudinal experiments on very young children with ASD are critically needed to better delineate modulations in brain patterns at the time of diagnosis, and how these alterations are influenced by therapeutic intervention. The present experiment is a first step towards that direction.

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Author details

1. Holger Franz Sperdin

   Developmental Imaging and Psychopathology Laboratory, Department of Psychiatry, University of Geneva, Geneva, Switzerland

   Contribution

   Conceptualization, Formal analysis, Investigation, Visualization, Methodology, Writing—original draft, Writing—review and editing

   Contributed equally with

   Ana Coito

   For correspondence

   holger.sperdin@unige.ch

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-3438-1572

2. Ana Coito

   Functional Brain Mapping Laboratory, Department of Fundamental Neurosciences, University of Geneva, Geneva, Switzerland

   Contribution

   Conceptualization, Software, Formal analysis, Validation, Investigation, Visualization, Methodology, Writing—original draft, Writing—review and editing

   Contributed equally with

   Holger Franz Sperdin

   Competing interests

   No competing interests declared

3. Nada Kojovic

   Developmental Imaging and Psychopathology Laboratory, Department of Psychiatry, University of Geneva, Geneva, Switzerland

   Contribution

   Software, Formal analysis, Investigation, Visualization

   Competing interests

   No competing interests declared

4. Tonia Anahi Rihs

   Functional Brain Mapping Laboratory, Department of Fundamental Neurosciences, University of Geneva, Geneva, Switzerland

   Contribution

   Conceptualization, Investigation, Writing—review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-6290-7439

5. Reem Kais Jan

   Functional Brain Mapping Laboratory, Department of Fundamental Neurosciences, University of Geneva, Geneva, Switzerland

   Contribution

   Investigation, Writing—review and editing

   Competing interests

   No competing interests declared

6. Martina Franchini

   Developmental Imaging and Psychopathology Laboratory, Department of Psychiatry, University of Geneva, Geneva, Switzerland

   Contribution

   Investigation, Writing—review and editing

   Competing interests

   No competing interests declared

7. Gijs Plomp

   Department of Psychology, University of Fribourg, Fribourg, Switzerland

   Contribution

   Software, Methodology, Writing—review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-9883-3371

8. Serge Vulliemoz

   EEG and Epilepsy Unit, Neurology, University Hospitals of Geneva, Geneva, Switzerland

   Contribution

   Supervision, Funding acquisition, Writing—review and editing

   Competing interests

   No competing interests declared

9. Stephan Eliez

   Developmental Imaging and Psychopathology Laboratory, Department of Psychiatry, University of Geneva, Geneva, Switzerland

   Contribution

   Resources, Supervision, Funding acquisition

   Competing interests

   No competing interests declared

10. Christoph Martin Michel

    Functional Brain Mapping Laboratory, Department of Fundamental Neurosciences, University of Geneva, Geneva, Switzerland

    Contribution

    Conceptualization, Resources, Software, Supervision, Funding acquisition, Methodology, Project administration, Writing—review and editing

    Competing interests

    No competing interests declared

11. Marie Schaer

    Developmental Imaging and Psychopathology Laboratory, Department of Psychiatry, University of Geneva, Geneva, Switzerland

    Contribution

    Conceptualization, Resources, Data curation, Software, Supervision, Funding acquisition, Investigation, Methodology, Project administration, Writing—review and editing

    Competing interests

    No competing interests declared

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