Facial expressions are critical to interpersonal communication and offer a nuanced, dynamic, and context-dependent insight into internal mental states. Humans use facial affect to infer personality, intentions, and emotions, and it is an important component of the clinical assessment of psychiatric illness. For these reasons, there has been significant interest in the objective analysis of facial affect (Naumann et al., 2009; Schmidt and Cohn, 2001; Ambadar et al., 2005). However, decisive techniques for quantifying facial affect under naturalistic conditions remain elusive.

A traditional approach is to count the occurrences of a discrete list of ‘universal basic emotions’ (Ekman, 1992). While the most commonly used system designates six basic emotions, there is disagreement about the number and nature of such affective ‘natural forms’ (Ortony, 2022; Keltner et al., 2019). Quantifying facial affect using the Facial Action Coding System (FACS) has become the dominant technique to operationalise facial expressions (Ekman et al., 1978). Action units, each corresponding to an anatomical facial muscle group, are rated on a quantitative scale. Traditional emotion labels are associated with the co-occurrence of a specific set of action units – for example, a ‘happy’ facial expression corresponds to action units ‘Cheek Raiser’ and ‘Lip Corner Puller’ (Friesen and Ekman, 1983). However, due to the time-intensive nature of manually coding every frame in a video, FACS has traditionally been applied to the analysis of static pictures rather than videos of human faces.

Recent developments in machine learning can assist the identification of basic emotions and facial action units from images and videos of human faces. Feature extraction for images include local textures (Feng, 2004; Kumar et al., 2016) and 3D geometry (Tian et al., 2001; Ghimire and Lee, 2013), while video analysis benefits from temporal features such as optical flow (Yeasin et al., 2006). Supervised learning algorithms classifying basic facial expressions based on feature values have achieved accuracies of 75–98% when benchmarked against manually coded datasets of both posed and spontaneous expressions (Ekundayo and Viriri, 2019). Endeavours such as the Facial Expression Recognition and Analysis challenge (Valstar et al., 2015) have improved cross-domain generalisability by encouraging the field to adopt common performance metrics.

Videos of faces can now be transformed into action unit time series which capture the rich temporal dynamics of facial expressions (Tian et al., 2001). This is important because human faces express emotional states dynamically – such as in a giggle or a sob. However, the rich potential of these temporal dynamics has not yet been fully exploited in the psychological and behavioural sciences. For example, some psychological studies and databases have asked responders to pose discrete emotions such as happiness or sadness (Lucey et al., 2019). This strategy suits the needs of a classic factorial experimental design but fails to produce the natural dynamics of real-world facial expressions. To evoke dynamic emotion, clinical interviews have been used (Darzi et al., 2019), or participants have been asked to narrate an emotive story or been shown emotive pictures rather than videos (Lang et al., 2008). Such pictures can be grouped into distinct categories and presented repetitively in a trial structure, but their ecological validity is unclear. Consequently, there is an expanding interest in using participants’ facial responses to naturalistic video stimuli (Kollias et al., 2019; Mavadati et al., 2019; Soleymani et al., 2012). These are more ecologically valid, have greater test-retest reliability than interviews, evoke stronger facial expressions than static pictures, and produce stronger cortical responses during functional neuroimaging (Ambadar et al., 2005; Sonkusare et al., 2019; Schultz and Pilz, 2009). However, interpreting participants’ facial expressions resulting from naturalistic stimulus viewing poses challenges, because each time point is unique. There is currently no obvious way to parse the stimulus video into discrete temporal segments. Naïve attempts at dimensionality reduction – for example, averaging action unit activations across time – omit temporal dynamics and so fail to capture the complexity of natural responses.

Disturbances in facial affect occur across a range of mental health disorders, including major depressive disorder, schizophrenia, and dementia. Capturing the nuances of facial affect is a crucial skill in clinical psychiatry but in the absence of quantitative tests this remains dependent on clinical opinion. Supervised learning has shown promise in distinguishing people with major depression from controls, using input features such as facial action units coded manually (Cohn et al., 2009) or automatically (Gavrilescu and Vizireanu, 2019), or model-agnostic representations of facial movements such as the ‘Bag of Words’ approach (Dibeklioğlu et al., 2015; Bhatia et al., 2017a). Studies documenting action unit occurrence during the course of a naturalistic stimulus (Renneberg et al., 2005), a short speech (Trémeau et al., 2005), or a clinical interview Girard et al., 2014 have demonstrated that depression is associated with reduced frequency of emotional expressions, particularly expressions with positive valence. Unfortunately, by averaging action unit occurrence over time, these methods poorly operationalise the clinician’s gestalt sense of affective reactivity, which derive from a patient’s facial responses across a range of contexts.

Here, we present a novel pipeline for processing facial expression data recorded while participants view a dynamic naturalistic stimulus. The approach is data-driven, eschewing the need to preselect emotion categories or segments of the stimulus video. We derive a time-frequency representation of facial movement information, on the basis that facial movements in vivo are fundamentally dynamic and multiscale. These time-frequency representations are then divided into discrete packets with a hidden Markov model (HMM), a method for inferring hidden states and their transitions from noisy observations. We find dynamic patterns of facial behaviour which are expressed sequentially and localised to specific action units and frequency bands. These patterns are context-dependent, consistent across participants, and correspond to intuitive concepts such as giggling and grimacing. We first demonstrate the validity of this approach on an open-source dataset of facial responses of healthy adults watching naturalistic stimuli. We then test this approach on facial videos of participants with melancholic depression, a severe mood disorder characterised by psychomotor changes (Parker and Hadzi Pavlovic, 1996). The time-frequency representation improves accuracy in classifying patients from healthy controls. However, unlike end-to-end deep learning approaches, our use of spectral features of facial action units facilitates easy interpretation while also exploiting prior knowledge to reduce the need for very large training datasets. Dynamic facial patterns reveal specific changes in melancholia, including reduced facial activity in response to emotional stimuli, anomalous facial responses inconsistent with the affective context, and a tendency to get ‘stuck’ in negatively valenced states.

We first analysed dynamic facial expressions from video recordings of 27 participants viewing short emotive clips of 4 minutes duration, covering a breadth of basic emotions (the Denver Intensity of Spontaneous Facial Action [DISFA] dataset [Mavadati et al., 2019] detailed in Supplementary file 1). Frame-by-frame action unit activations were extracted with OpenFace software (Baltrusaitis et al., 2018) (see Supplementary file 1 for action unit descriptions, Figure 1—figure supplement 1 for group mean time series).

From these data, we used the continuous wavelet transform to extract a time-frequency representation of individual action unit time series in each participant. To test whether this time-frequency representation captures high-frequency dynamic content, we first compared the group average of these individual time-frequency representations with the time-frequency representation of the group mean time series. We selected the activity of action unit 12 ‘Lip Corner Puller’ during a positive valence video clip (a ‘talking dog’), as this action unit is conventionally associated with happy affect, and its high-frequency activity denotes smiling or laughing. Compared to the group mean time series, the time series of individuals had significantly greater amplitude (Figure 1b), particularly at higher frequencies (Figure 1f). This demonstrates that the time-frequency representations of individual participants capture high-frequency dynamics that are obscured by characterising group-averaged time courses. This is because stimulus-evoked facial action unit responses have asynchronous alignment across participants, hence cancelling when superimposed. This problem is avoided in the group-level time-frequency representation, whereby the amplitude is first extracted at the individual level, prior to group averaging. Comparable results occurred in all action units (Figure 1—figure supplement 2).

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Having shown how the time-frequency representation captures dynamic content, we next sought to quantify the joint dynamics of facial action units. To this end, an HMM was inferred from the time course of all facial action units’ time-frequency representations. An HMM infers a set of distinct states from noisy observations, with each state expressed sequentially in time according to state-to-state transition probabilities. Each state has a distinct mapping onto the input space, here the space of frequency bands and action units (see Figure 2 for a visual overview of the entire pipeline, Figure 3a for the HMM states derived from the DISFA dataset).

Figure 2

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Examples of participants in each state are provided Figure 3—videos 1–19. Their occurrence corresponded strongly with annotated video clip valence (Figure 3b). We found that inferred state sequences had high between-subject consistency, exceeding chance level across the vast majority of time points and reaching 93% during specific movie events (Figure 3d). States were frequency-localised and comprised intuitive combinations of action units which reflected not only distinct emotion categories as defined in previous literature (Friesen and Ekman, 1983), but also stimulus properties such as mixed emotions. State transition probabilities appeared clustered by valence rather than frequency, such that frequent transitions between low- and high-frequency oscillations of the same facial action units were more likely than transitions between different emotions (Figure 3e).

• States 1 and 2 were active during stimuli annotated as ‘happy’. They activated two action units typically associated with happiness, action unit 6 ‘Cheek Raiser’ and 12 ‘Lip Corner Puller’, but also action unit 25 ‘Lips Part’. State 2 likely represents laughing or giggling as it encompassed high-frequency oscillations in positive valence action units, in comparison to the low-frequency content of state 1.

• States 3 and 4 were active during videos evoking fear and disgust – for example of a man eating a beetle larva. They encompassed mixtures of action units conventionally implicated in disgust and fear, at low- and high-frequency bands, respectively. State 3 recruited action units 4 ‘Brow Lowerer’ and 9 ‘Nose Wrinkler’, while state 4 involved these action units as well as action units 15 ‘Lip Corner Depressor’, 17 ‘Chin Raiser’, and 20 ‘Lip Stretcher’.

• States 5 and 6 occurred predominantly during negatively valenced clips, and deactivated oscillatory activity in most action units, with sparing of action units typically associated with sadness, 4 ‘Brow Lowerer’ and 15 ‘Lip Corner Depressor’.

Group time courses in melancholia

Facial action unit time courses showed clear group differences (see Figure 4 for action units typically implicated in expressing happiness and sadness, and Figure 4—figure supplement 3 for all action units). For each action unit in each participant, we calculated the median action unit activation across each stimulus video. These were compared with a three-way ANOVA, with factors for clinical group, stimulus, and the facial valence. We considered two stimulus videos, one with positive and one with negative valence, and two facial valence states, happiness and sadness, calculated as sums of positively and negatively valenced action unit activations, respectively (Friesen and Ekman, 1983). A significant three-way interaction was found between clinical group, stimulus, and facial valence (p=0.003). Post hoc comparisons with Tukey’s honestly significant difference criterion (Figure 4—figure supplement 1) quantified that during stand-up comedy, participants with melancholia had reduced activation of action unit 12 ‘Lip Corner Puller’ (p<0.0001) and increased activation of action unit 4 ‘Brow Lowerer’ (p<0.0001). Interestingly, facial responses of participants with melancholia during stand-up comedy were similar to those of controls during the sad movie (p>0.05 for both action units). Results were unchanged when using the weather report stimulus instead of the stand-up comedy (p=0.005 for three-way interaction, see Figure 4—figure supplement 2 for post hoc comparisons).

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To move away from individual action units, we next extracted the first principal component across all action units. The time course of this composite component closely followed joke punch lines during stand-up comedy (Figure 4b). This responsivity of this component to movie events was substantially diminished in the melancholia cohort.

Time-frequency representation in melancholia

Time-frequency representations were calculated for all action units in all participants. For each action unit, the mean time-frequency representation for the control group was subtracted from the participants with melancholia (see Figure 5—figure supplement 1 for the mean of the controls). Significant group differences (p<0.05) were found by comparison to a null distribution composed of 1000 resampled surrogate datasets (see Materials and methods). Participants with melancholia had a complex pattern of reduced activity encompassing a broad range of frequencies (Figure 5a). The most prominent differences were in positive valence action units during positive valence stimuli, but significant reductions were seen in most action units. Differences in high-frequency bands occurred during specific movie events such as jokes (Figure 5b). There were sporadic instances of increased activity in melancholia participants during the sad movie involving mainly action units 15 ‘Lip Corner Depressor’ and 20 ‘Lip Stretcher’.

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We next pursued whether the additional time-frequency information would improve the classification accuracy of differentiating participants with melancholia from controls. A support vector machine, using as inputs the mean action unit activation for each stimulus video, achieved 63% accuracy with five-fold cross-validation. In contrast, using as inputs the mean time-frequency amplitude in discrete frequency bands within 0–5 Hz improved average cross-validation accuracy to 71% (p<0.001 for difference between models). As a control for the additional number of input features, we tested a third set of models which naively modelled temporal dynamics using mean action unit activations within shorter time blocks. These models had 63–64% accuracy despite having a greater number of input features than the time-frequency representation (Supplementary file 1).

Sequential affective states in melancholia

Inverting a HMM from the time-frequency representations of facial action units yielded the sequential expression of eight states across participants (Figure 6).

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• States 1 and 2 activated positive valence action units, each in distinct frequency bands, and were dominant through the stand-up comedy for most participants (Figure 6b). State 2 comprised high-frequency oscillations in positive valence action units, corresponding to laughing or giggling.

• The sad movie was associated with early involvement of state 3, which deactivated high-frequency activity, followed by states 4 and 5, which also deactivated oscillatory activity, but with more specificity for lower frequencies and positive valence action units.

• State 6 comprised action units 4 ‘Brow Lowerer’, 9 ‘Nose Wrinkler’, 17 ‘Chin Raiser’, and 23 ‘Lip Tightener’, traditionally associated with anger, disgust, or concern. State 7 can be associated with ‘gasping’, with very high-frequency activation of most mouth-associated action units including 25 ‘Lips Part’. These states occurred sporadically through the weather report.

• State 8 predominantly activated action unit 1 ‘Inner Brow Raiser’, commonly associated with negative valence.

The temporal sequence of the most common state was similar across groups (Figure 6c), but the between-subjects consistency was markedly reduced in the melancholic participants during both funny videos (Figure 6d). Some participants with melancholia – for example participants 2 and 3 (Figure 6b) – had highly anomalous state sequences compared to other participants.

Fractional occupancy – the proportion of time spent by participants in each state – was significantly different between groups for the positive valence states – state 1 (Melancholia < Controls, p_FDR = 0.03) and state 2 (Melancholia < Controls, p_FDR = 0.004) – as well as for negatively valenced state 8 (Melancholia > Controls p_FDR = 0.03). We then asked whether group differences in the time spent in each state were attributable to changes in the likelihood of switching in to, or out of, specific facial states. Participants with melancholia were significantly less likely to switch from a low-frequency positive valence state (1, smiling) to high-frequency positive valence oscillations (state 2, giggling), but were more likely to switch to states associated with any other emotion (states 4, 5, 6, 7, and 8). From the high-frequency positive valence state, they were more likely to switch to the deactivating ‘ennui’ state 4 (all p_FDR < 0.05).

Facial expressions played a crucial role in the evolution of social intelligence in primates (Schmidt and Cohn, 2001) and continue to mediate human interactions. Observations of facial affect, its range, and reactivity play a central role in clinical settings. Quantitative analysis of facial expression has accelerated of late, driven by methods to automatically factorise expressions into action units (Ekman et al., 1978) and the availability of large datasets of posed emotions (Lucey et al., 2019). The dynamics of facial expression mediate emotional reciprocity, but have received less attention (Ambadar et al., 2005). Naturalistic stimuli offer distinct advantages to affective research for their ability to evoke these dynamic responses (Dhall et al., 2012), but their incompressibility has made analysis problematic. By leveraging techniques in computer vision, we developed a pipeline to characterise facial dynamics during naturalistic video stimuli. Analysis of healthy adults watching emotionally salient videos showed that facial expression dynamics can be captured by a small number of spatiotemporal states. These states co-activate facial muscle groups with a distinct spectral fingerprint, and transition dynamically with the emotional context. Application of this approach to melancholia showed that the clinical gestalt of facial non-reactivity in melancholia (Parker, 2007) can be objectively identified not just with restrictions in spectral content, but also with anomalous facial responses, more frequent occurrence of an ennui affect, and more frequent state switching from transiently positive facial expressions to neutral and negative states. This approach provides a unique perspective on how facial affect is generated by the interplay between inner affective states and the sensorium.

Our pipeline first comprises automatic action unit extraction, then spectral wavelet-based analysis of the ensuing feature dynamics. Wavelet energy at a given time corresponds to the occurrence of a specific facial ‘event’, while energy in a given frequency reflects the associated facial dynamics, like laughing. Unlike temporal averaging methods, which require an arbitrary timescale, wavelets cover a range of timescales. The spectral approach also allows participant facial responses to be pooled, without the limitations of averaging responses whose phases are misaligned. We then inferred an HMM, identifying spatially and spectrally resolved modes of dynamic facial activity which occur sequentially with high consistency across participants viewing the same stimulus. States transitions aligned with intuitive notions of affective transitions. For example, the common transition between the low- and high-frequency positive valence state reflected transitions between smiling and laughing.

Our method builds on the Emotion Facial Action Coding System (EMFACS) (Friesen and Ekman, 1983), where each canonical emotion label (happy, angry, etc.) is defined on the basis of a sparse set of minimally necessary action units. The sparsity of this coding allows manual raters to find the minimal necessary combinations of action units in a facial video to reflect an emotion label, but may not include all action units that are involved in each affective state. Affective states inferred from our HMM not only reflected prototypical action unit combinations from EMFACS, but also provide a richer mapping across a broader range of action units. For example, while happiness has been previously associated with just two action units, ‘Cheek Raiser’ and ‘Lip Corner Puller’, such sparse activations are rare, particularly during intense emotional displays. We demonstrated that laughing during stand-up comedy activated eyebrow-related action units, some of which are traditionally associated with sadness. Conversely, negatively valenced stimuli dampened facial movements, with a relative sparing of those action units typically associated with sadness.

Ensembles of HMMs have previously been used to improve emotion classification accuracy when benchmarked against manually coded datasets. In these studies, one HMM models the temporal dynamics of one action unit (Jiang et al., 2014; Koelstra et al., 2010) or one universal basic emotion (Yeasin et al., 2006; Sandbach et al., 2012), with HMM states corresponding to expression onset/offset. Given a video frame, the HMM with the greatest evidence determines the decoded expression. Nested HMMs have also been employed, with a second-level HMM predicted transitions between the basic emotions (Cohen et al., 2012). In contrast, the present method uses a single HMM to describe facial expressions without prior emotion categories, capturing the dynamic co-occurrence of facial actions that together comprise distinct affective states. By taking the spectral activity of action units as input features into the HMM, our approach uniquely captures the spatiotemporal texture of naturally occurring facial affect. This enables, for example, the disambiguation of a smile from a giggle. The importance of the spectral characterisation is highlighted by our finding that in melancholia, smile states were more likely to transition to ennui, and less likely to the laughter state. Our use of dynamic spectra as inputs into an HMM is similar to their recent use in neuroimaging research (Baker et al., 2014). Using the raw time series is also possible – hence additionally capturing phase relationships, although this comes with an additional computational burden and reduced interpretability of states (Vidaurre et al., 2018).

Dynamic facial patterns were influenced by the affective properties of the stimulus video. For the DISFA dataset, the HMM inferred two disgust-associated states, in low- and high-frequency bands, respectively. These states occurred predominantly during two disgusting video clips. For the melancholia dataset, the inferred HMM states over-represented happiness and sadness, and under-represented disgust. This is ostensibly because the stimulus had prominent positively and negatively valenced sections without disgusting content. The co-occurrence of the states and the state transitions across participants speaks to the influence of the video content on affective responses and hence, more broadly, the dynamic exchange between facial affect and the social environment.

We found that participants with melancholia exhibited broad reductions in facial activity, as well as specific reductions in high-frequency activity in response to joke punchlines, reflecting the clinical gestalt of impaired affective reactivity to positively valenced events (Parker and Hadzi Pavlovic, 1996). Unlike previous reports which highlighted reduced reactivity to negative stimuli in depression (Bylsma et al., 2008), we did not find significant group differences for negative expression reactivity. Viewing affect as a dynamic process provided two further insights into facial responses in melancholia. First, decreased between-subject consistency and more anomalous facial responses suggest that their facial activity is less likely to be driven by a common external stimulus. Ambiguous facial responses are also seen in schizophrenia (Hamm et al., 2014), suggesting the possibility of a common underlying mechanism with melancholia. Second, participants with melancholia were less likely to enter high-frequency positive valence states like laughing, and once there, transitioned out quickly to the ‘ennui’ state. This reflects the clinical impression that positive mood states persist in healthy controls, but such states are fleeting in those with melancholia, who tend to get ‘stuck’ in negative mood states instead. The results are commensurate with the proposal that depressed states relate to persistent firing in non-reward functional areas mediated by attractor dynamics (Rolls, 2016). Additionally, these findings accord with neurobiological models of melancholia whereby dysfunctional cortical-basal ganglia circuitry underlie the disturbances in volition and psychomotor activity that characterise the disorder (Parker and Hadzi Pavlovic, 1996). More generally, the notion of affect as a sequence of spatiotemporal states aligns with the proposal that instabilities in brain network activity generate adaptive fluctuations in mood and affect, with these being either over- or under-damped in affective disorders (Perry et al., 2019). Our paradigm also raises clinical questions predicated on dynamics – for example, do biological or psychological treatments for melancholia work by increasing the probability of entering positive affective states, or reducing the probability of exiting such states?

Several caveats bear mention. First, it is well known that cross-domain generalisability in facial affect detection trails behind within-domain performance (Ruiz et al., 2017; Li et al., 2017) (see Cohn et al., 2019, for a review). However, the spectral transform method is dependent on sensitivity to changes in AU intensity, rather than on the accuracy of predicted AU occurrence. Cross-domain generalisability may be closer to adequate for AU intensity than for AU occurrence, because appearance features such as the nasiolabial furrow depth vary continuously with AU intensity (Cohn et al., 2019). Conversely, AU occurrence detection may depend on specific feature value thresholds that vary in different datasets due to illumination, face orientation, gender, and age. We also have greater confidence in OpenFace’s sensitivity to changes in AU intensity, because its ability to locate landmark points has been validated on a truly out-of-sample dataset (300-VW dataset) (Baltrusaitis et al., 2018). Second, a small number of participants with constant zero activation in one or more action units were excluded from analysis, because this produces an ill-defined spectral transform. Excluded participants, of whom one was a control and four had melancholia, may have had the greatest impairments in facial affect. This issue could be addressed with a lower detectable limit of action unit activation. Third, time-frequency maps were standardised in mean and variance before HMM inference. This ensures that states occur sequentially across time, but reduces the differences in state sequences across groups. Omitting this standardisation step yields states that are biased towards group differences rather than temporal differences (see Figure 6—figure supplement 1). Future work could consider methods that are less susceptible to this trade-off. Fourth, the generalisability of our results in the clinical group is limited by sample size, and would benefit from independent replication before clinical applications are considered. Fifth, the modest inter-rater reliability of some psychiatric diagnoses (Aboraya et al., 2006) raises questions about the extent to which diagnoses of melancholic depression can be considered as ‘ground truth’. However, we have previously demonstrated changes in functional brain connectivity in this cohort (Guo et al., 2016), highlighting the neurobiological difference between these groups. Sixth, the present pipeline focused on oscillatory magnitudes and discarded phase information from the wavelet transform. Future work could incorporate phase information to quantify how facial responses synchronize with stimulus properties. Finally, the utility of our approach is likely to be improved by multimodal fusion of facial, head pose, vocal, and body language behaviour, each of which independently improve classification (Bhatia et al., 2017b; Alghowinem et al., 2006b; Alghowinem et al., 2006a; Joshi et al., 2006).

Human emotion and affect are inherently dynamic. Our work demonstrates that momentary affective responses, such as laughing or grimacing, traditionally viewed from a qualitative standpoint, can be understood within a quantitative framework. These tools are cheap, automatable, and could be used within a smartphone operating system to complement the brief assessment of facial affect during a clinical encounter. Potential translational applications include screening for mental health disorders or monitoring clinical progress. Quantifying dynamic features of facial affect could also assist in subtyping the phenomenology of reduced expressivity, to distinguish between psychomotor retardation in melancholic depression, emotional incongruence, and affective blunting in schizophrenia, the masked facies of Parkinson’s disease, or apathy in dementia. Steps towards this translation will require evaluation of its acceptability and utility in clinical practice.

The DISFA dataset is publically available at http://mohammadmahoor.com/disfa/, and can be accessed by application at http://mohammadmahoor.com/disfa-contact-form/. The melancholia dataset is not publically available due to ethical and privacy considerations for patients, and because the original ethics approval does not permit sharing this data.

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Author details

1. Jayson Jeganathan

   1. School of Psychology, College of Engineering, Science and the Environment, University of Newcastle, Newcastle, Australia
   2. Hunter Medical Research Institute, Newcastle, Australia

   Contribution

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

   For correspondence

   jayson.jeganathan@gmail.com

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-4175-918X

2. Megan Campbell

   1. School of Psychology, College of Engineering, Science and the Environment, University of Newcastle, Newcastle, Australia
   2. Hunter Medical Research Institute, Newcastle, Australia

   Contribution

   Conceptualization, Supervision, Methodology, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-4051-1529

3. Matthew Hyett

   School of Psychological Sciences, University of Western Australia, Perth, Australia

   Contribution

   Resources, Data curation, Methodology, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

4. Gordon Parker

   School of Psychiatry, University of New South Wales, Kensington, Australia

   Contribution

   Resources, Data curation, Funding acquisition, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

5. Michael Breakspear

   1. School of Psychology, College of Engineering, Science and the Environment, University of Newcastle, Newcastle, Australia
   2. Hunter Medical Research Institute, Newcastle, Australia
   3. School of Medicine and Public Health, College of Medicine, Health and Wellbeing, University of Newcastle, Newcastle, Australia

   Contribution

   Conceptualization, Resources, Supervision, Methodology, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-4943-3969

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