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Social deficits and dysregulations in dopaminergic midbrain-striato-frontal circuits represent transdiagnostic symptoms across psychiatric disorders. Animal models suggest that interactions between the dopamine (DA) and renin-angiotensin system (RAS) may modulate learning and reward-related processes. The present study therefore examined the behavioral and neural effects of the Angiotensin II type 1 receptor (AT1R) antagonist losartan on social reward and punishment processing in humans. A preregistered randomized double-blind placebo-controlled between-subject pharmacological design was combined with a social incentive delay (SID) functional MRI (fMRI) paradigm during which subjects could avoid social punishment or gain social reward. Healthy volunteers received a single-dose of losartan (50 mg,n= 43, female = 17) or placebo (n= 44, female = 20). We evaluated reaction times (RTs) and emotional ratings as behavioral and activation and functional connectivity as neural outcomes. Relative to placebo, losartan modulated the reaction time and arousal differences between social punishment and social reward. On the neural level the losartan-enhanced motivational salience of social rewards was accompanied by stronger ventral striatum-prefrontal connectivity during reward anticipation. Losartan increased the reward-neutral difference in the ventral tegmental area (VTA) and attenuated VTA associated connectivity with the bilateral insula in response to punishment during the outcome phase. Thus, losartan modulated approach-avoidance motivation and emotional salience during social punishment versus social reward via modulating distinct core nodes of the midbrain-striato-frontal circuits. The findings document a modulatory role of the renin-angiotensin system in these circuits and associated social processes, suggesting a promising treatment target to alleviate social dysregulations.SIGNIFICANCE STATEMENTSocial deficits and anhedonia characterize several mental disorders and have been linked to the midbrain-striato-frontal circuits of the brain. Based on initial findings from animal models we here combine the pharmacological blockade of the Angiotensin II type 1 receptor (AT1R) via losartan with functional MRI (fMRI) to demonstrate that AT1R blockade enhances the motivational salience of social rewards and attenuates the negative impact of social punishment via modulating the communication in the midbrain-striato-frontal circuits in humans. The findings demonstrate for the first time an important role of the AT1R in social reward processing in humans and render the AT1R as promising novel treatment target for social and motivational deficits in mental disorders.

Background The neuropeptide oxytocin (OXT) modulates social cognition by increasing attention to social cues and may have therapeutic potential for impaired social attention in conditions such as autism spectrum disorder. Intranasal administration of OXT is widely used to examine the drug’s functional effects in both adults and children and is assumed to enter the brain directly via this route. However, OXT can also influence brain function through increased blood concentrations, and we have recently shown that orally (lingual) administered OXT also modulates neural responses to emotional faces and may be better tolerated for therapeutic use. Here, we examine whether 24 IU OXT administered orally can facilitate social attention. Methods In a randomized, placebo-controlled pharmacologic study, we used a validated emotional antisaccade eye-tracking paradigm to explore the effects of oral OXT on bottom-up and top-down attention processing in 80 healthy male participants. Results Our findings showed that in terms of top-down attention, oral OXT increased errors for both social (angry, fearful, happy, sad, and neutral emotion faces) and nonsocial stimuli (oval shapes) in the antisaccade condition but increased response latencies only in the social condition. It also significantly reduced post-task state anxiety, but this reduction was not correlated with task performance. A comparison with our previous intranasal OXT study using the same task revealed that both routes have a similar effect on increasing antisaccade errors and response latencies and on reducing state anxiety. Conclusions Overall, our findings suggest that oral administration of OXT produces similar effects on top-down social attention control and anxiety to intranasal administration and may therefore have therapeutic utility.

There is considerable interest in the significance of structural and functional connections between the two brain hemispheres in terms of both normal function and in relation to psychiatric disorders. In recent years, many studies have used voxel mirrored homotopic connectivity analysis of resting state data to investigate the importance of connectivity between homotopic regions in the brain hemispheres in a range of neuropsychiatric disorders. The current review summarizes findings from these voxel mirrored homotopic connectivity studies in individuals with autism spectrum disorder, addiction, attention deficit hyperactivity disorder, anxiety and depression disorders, and schizophrenia, as well as disorders such as Alzheimer's disease, mild cognitive impairment, epilepsy, and insomnia. Overall, other than attention deficit hyperactivity disorder, studies across psychiatric disorders report decreased homotopic resting state functional connectivity in the default mode, attention, salience, sensorimotor, social cognition, visual recognition, primary visual processing, and reward networks, which are often associated with symptom severity and/or illness onset/duration. Decreased homotopic resting state functional connectivity may therefore represent a transdiagnostic marker for general psychopathology. In terms of disorder specificity, the extensive decreases in homotopic resting state functional connectivity in autism differ markedly from attention deficit hyperactivity disorder, despite both occurring during early childhood and showing extensive co-morbidity. A pattern of more posterior than anterior regions showing reductions in schizophrenia is also distinctive. Going forward, more studies are needed to elucidate the functions of these homotopic functional connections in both health and disorder and focusing on associations with general psychopathology, and not only on disorder specific symptoms.

Generalized Anxiety Disorder (GAD) and Major Depressive Disorder (MDD) are both characterized by cognitive and social impairments. Determining disorder-specific neurobiological alterations in GAD and MDD by means of functional magnetic resonance imaging (fMRI) may promote determination of precise diagnostic markers.This study aimed to examine disorder-specific behavioral and neural alterations at the intersection of social and cognitive processing in treatment-naïve first-episode GAD (n = 35) and MDD (n = 37) patients compared to healthy controls (n = 35) by employing a social-emotional n-back fMRI paradigm.No behavioral differences between patients and healthy controls were observed. However, GAD patients exhibited decreased bilateral dorsomedial prefrontal cortex (dmPFC) engagement during the 0-back condition yet increased dmPFC engagement during the 1-back condition compared to MDD and healthy participants. In contrast, MDD patients exhibited increased dmPFC-insula coupling during 0-back, yet decreased coupling during 1-back, compared to GAD and healthy participants. Dimensional symptom-load analysis confirmed that increased dmPFC-insula connectivity during 0-back was positively associated with depressive symptom load.The moderate sample size in the present study did not allow us to further explore gender differences. In addition, some patients exhibited GAD and MDD comorbidity according to the M.I.N.I. interview. Finally, the paradigm we used did not allow to further disentangle emotion-specific effects on working memory.These findings suggest that the dmPFC engaged in integrating affective and cognitive components and self-other processing exhibits GAD-specific neurofunctional dysregulations whereas functional dmPFC communication with the insula, a region involved in salience processing, may represent an MDD-specific neurofunctional deficit.

In recent years ample studies have reported that intranasal administration of the neuropeptide oxytocin can facilitate social motivation and cognition in healthy and clinical populations. However, it is still unclear how effects are mediated since intranasally administered oxytocin can both directly enter the brain (nose to brain) and increase peripheral vascular concentrations (nose to blood). The relative functional contributions of these routes are not established and have received insufficient attention in the field. The current study used vasoconstrictor pretreatment to prevent intranasal oxytocin (24 IU) from increasing peripheral concentrations and measured effects on both resting-state neural (electroencephalography) and physiological responses (electrocardiogram, electrogastrogram and skin conductance). Results demonstrated that intranasal oxytocin alone produced robust and widespread increases of delta-beta cross-frequency coupling (CFC) from 30 min post-treatment but did not influence peripheral physiological measures. As predicted, vasoconstrictor pretreatment greatly reduced the normal increase in peripheral oxytocin concentrations and, importantly, abolished the majority of intranasal oxytocin effects on delta-beta CFC. Furthermore, time-dependent positive correlations were found between increases in plasma oxytocin concentrations and corresponding increases in delta-beta CFC following oxytocin treatment alone. Our findings suggest a critical role of peripheral vasculature-mediated routes on neural effects of exogenous oxytocin administration with important translational implications for its use as an intervention in psychiatric disorders.

Background The hypothalamic neuropeptide oxytocin (OXT) may exert anxiolytic and stress-reducing actions via modulatory effects on amygdala circuits. Animal models and initial findings in humans suggest that some of these effects are mediated by interactions with other neurotransmitter systems, in particular the serotonin (5-HT) system. Against this background, the present pharmacological resting-state functional magnetic resonance imaging study aimed to determine whether effects of OXT on stress-associated amygdala intrinsic networks are mediated by 5-HT. Methods We employed a randomized, placebo-controlled, double-blind parallel-group, pharmacological functional magnetic resonance imaging resting-state experiment with 4 treatment groups in n = 112 healthy male participants. Participants underwent a transient decrease in 5-HT signaling via acute tryptophan depletion (ATD) or a corresponding placebo-control protocol before the administration of intranasal OXT (24 IU) or placebo intranasal spray. Results OXT and 5-HT modulation exerted interactive effects on the coupling of the left amygdala with the ipsilateral hippocampus and adjacent midbrain. OXT increased intrinsic coupling in this pathway, whereas this effect of OXT was significantly attenuated during transiently decreased central serotonergic signaling induced via acute tryptophan depletion. In the absence of OXT or 5-HT modulation, this pathway showed a trend for an association with self-reported stress perception in everyday life. No interactive effects were observed for the right amygdala. Conclusions Together, the findings provide the first evidence, to our knowledge, that the effects of OXT on stress-associated amygdala-hippocampal-midbrain pathways are critically mediated by the 5-HT system in humans.

Serotonin (5-HT) has long been implicated in adaptive emotion regulation as well as the development and treatment of emotional dysregulations in mental disorders. Accumulating evidence suggests that a genetic vulnerability may render some individuals at a greater risk for the detrimental effects of transient variations in 5-HT signaling. The present study aimed to investigate whether individual variations in the Tryptophan hydroxylase 2 (TPH2) genetics influence susceptibility for behavioral and neural threat reactivity dysregulations during transiently decreased 5-HT signaling. To this end, interactive effects between TPH2 (rs4570625) genotype and acute tryptophan depletion (ATD) on reactivity towards angry, neutral and happy faces were examined in a within-subject placebo-controlled pharmacological fMRI trial (n = 51). An a priori genotype stratification approach of extreme groups (GG vs. TT) allowed balanced sampling. While no main effects of ATD on neural reactivity to threat-related stimuli and mood state were observed in the entire sample, accounting for TPH2 genotype revealed an ATD-induced increase in subjective anxious arousal in the GG but not the TT carriers. The effects were mirrored on the neural level, such that ATD specifically reduced ventromedial prefrontal cortex (vmPFC) reactivity towards threat-related stimuli in the GG carriers. Furthermore, the ATD-induced increase in subjective anxiety positively associated with the extent of ATD-induced changes in vmPFC activity in response to threat-related stimuli in GG carriers. Together the present findings suggest for the first time that individual variations in TPH2 genetics render individuals susceptible to the anxiogenic and neural effects of a transient decrease in 5-HT signaling.

Background Autism spectrum disorder (ASD) is associated with altered brain development, but it is unclear which specific structural changes may serve as potential diagnostic markers, particularly in young children at the age when symptoms become fully established. Furthermore, such brain markers need to meet the requirements of precision medicine and be accurate in aiding diagnosis at an individual rather than only a group level. Objective This study aimed to identify and model brain-wide differences in structural connectivity using diffusion tensor imaging (DTI) in young ASD and typically developing (TD) children. Methods A discovery cohort including 93 ASD and 26 TD children and two independent validation cohorts including 12 ASD and 9 TD children from three different cities in China were included. Brain-wide (294 regions) structural connectivity was measured using DTI (fractional anisotropy, FA) together with symptom severity and cognitive development. A connection matrix was constructed for each child for comparisons between ASD and TD groups. Pattern classification was performed on the discovery dataset and the resulting model was tested on the two independent validation datasets. Results Thirty-three structural connections showed increased FA in ASD compared to TD children and associated with both autistic symptom severity and impaired general cognitive development. The majority (29/33) involved the frontal lobe and comprised five different networks with functional relevance to default mode, motor control, social recognition, language and reward. Overall, classification achieved very high accuracy of 96.77% in the discovery dataset, and 91.67% and 88.89% in the two independent validation datasets. Conclusions Identified structural connectivity differences primarily involving the frontal cortex can very accurately distinguish novel individual ASD from TD children and may therefore represent a robust early brain biomarker which can address the requirements of precision medicine.

Introduction: Anxiety disorders are prevalent mental conditions characterized by exaggerated anxious arousal and threat reactivity. Animal and human studies suggest an anxiolytic potential of the neuropeptide oxytocin (OT), yet, while a clinical application will require chronic administration protocols, previous human studies have exclusively focused on single-dose (acute) intranasal OT effects. Objective: To facilitate the translation of the potential anxiolytic mechanism of OT into clinical application, we determined whether the anxiolytic effects of OT are maintained with repeated (chronic) administration or are influenced by dose frequency and trait anxiety. Methods: In a pre-registered double-blind randomized placebo-controlled pharmaco-fMRI trial the acute (single dose) as well as chronic effects of two different dose frequencies of OT (OT administered daily for 5 days or every other day) on emotional reactivity were assessed in n = 147 individuals with high versus low trait anxiety (ClinicalTrials.gov ID: NCT03085654). Results: OT produced valence, dose frequency, and trait anxiety-specific effects, such that the low-frequency (intermittent) chronic dosage specifically attenuated a neural reactivity increase in amygdala-insula-prefrontal circuits observed in the high anxious placebo-treated subjects in response to threatening but not positive stimuli. Conclusions: The present trial provides the first evidence that low-dose frequency chronic intranasal OT has the potential to alleviate exaggerated neural threat reactivity in subjects with elevated anxiety levels, suggesting a treatment potential for anxiety disorders.

The cerebral cortex is folded as gyri and sulci, which provide the foundation to unveil anatomo-functional relationship of brain. Previous studies have extensively demonstrated that gyri and sulci exhibit intrinsic functional difference, which is further supported by morphological, genetic, and structural evidences. Therefore, systematically investigating the gyro-sulcal (G-S) functional difference can help deeply understand the functional mechanism of brain. By integrating functional magnetic resonance imaging (fMRI) with advanced deep learning models, recent studies have unveiled the temporal difference in functional activity between gyri and sulci. However, the potential difference of functional connectivity, which represents functional dependency between gyri and sulci, is much unknown. Moreover, the regularity and variability of the G-S functional connectivity difference across multiple task domains remains to be explored. To address the two concerns, this study developed new anatomy-guided spatio-temporal graph convolutional networks (AG-STGCNs) to investigate the regularity and variability of functional connectivity differences between gyri and sulci across multiple task domains. Based on 830 subjects with seven different task-based and one resting state fMRI (rs-fMRI) datasets from the public Human Connectome Project (HCP), we consistently found that there are significant differences of functional connectivity between gyral and sulcal regions within task domains compared with resting state (RS). Furthermore, there is considerable variability of such functional connectivity and information flow between gyri and sulci across different task domains, which are correlated with individual cognitive behaviors. Our study helps better understand the functional segregation of gyri and sulci within task domains as well as the anatomo-functional-behavioral relationship of the human brain.

BackgroundOverarching conceptualizations propose that the complex social-emotional effects of oxytocin (OXT) in humans are partly mediated by interactions with other neurotransmitter systems. Recent animal models suggest that the anxiolytic effects of OXT are critically mediated by the serotonin (5-HT) system, yet direct evidence in humans is lacking.MethodsTo determine the role of 5-HT in OXT-induced attenuation of amygdala threat reactivity and sensitization/ desensitization, we conducted a parallel-group randomized placebo-controlled double-blind experiment during which n = 121 healthy subjects underwent a transient decrease in 5-HT signaling via acute tryptophan depletion (ATD, TRYP-) or the corresponding placebo-control protocols before the administration of intranasal OXT or placebo intranasal spray, respectively. Mean and repetition-dependent changes in threat-specific amygdala reactivity towards threatening stimuli (angry faces) as assessed by fMRI served as the primary outcome.ResultsNo treatment main or interaction effects on amygdala threat reactivity were observed, yet OXT switched bilateral amygdala threat sensitization to desensitization and this effect was significantly attenuated during decreased central 5-HT signaling via pretreatment with TRYP-.ConclusionsThe present findings provide the first evidence for a role of OXT in threat-specific amygdala desensitization in humans and suggest that these effects are critically mediated by the 5-HT system. OXT may have a therapeutic potential to facilitate amygdala desensitization and adjunct up-regulation of 5-HT neurotransmission may facilitate OXT’s anxiolytic potential.The trial was preregistered on clinicaltrials.gov (https://clinicaltrials.gov/ct2/show/NCT03426176, ID NCT03426176)

Touch represents one of our most important senses throughout life and particularly in the context of our social and emotional experiences. In this review, we draw on research on touch processing from both animal models and humans. Firstly, we briefly describe the cutaneous touch receptors and neural processing of both affective and discriminative touch. We then outline how our sense of touch develops and summarize increasing evidence demonstrating how essential early tactile stimulation is for the development of brain and behavior, with a particular focus on effects of tactile stimulation in infant animals and pediatric massage and Kangaroo care in human infants. Next, the potential mechanisms whereby early tactile stimulation influences both brain and behavioral development are discussed, focusing on its ability to promote neural plasticity changes and brain interhemispheric communication, development of social behavior and bonding, and reward sensitivity through modulation of growth factor, oxytocin, and opioid signaling. Finally, we consider the implications of evidence for atypical responses to touch in neurodevelopmental disorders such as autism spectrum disorder and discuss existing evidence and future priorities for establishing potential beneficial effects of interventions using massage or pharmacological treatments targeting oxytocin or other neurochemical systems.

This study investigated heterogeneity in language skills of children with autism and their relationship with different autistic social subtypes. Data from 90 autistic and 30 typically developing children were analyzed. Results showed that autistic social subtypes varied in language skill problems (aloof passive active-but-odd). There was a negative association between aloof dimension scores and language performance but positive for the active-but-odd dimension and no association in the passive one. Moreover, aloof dimension score was the main contributor to language performance. A receiver operating characteristic analysis suggested language vocabulary as an additional component in differentiating autistic social subtypes. These findings demonstrate that variations in language skills in autistic children provide additional information for discriminating their social subtype.

Non-invasive transcutaneous auricular vagus nerve stimulation (taVNS) as a newly developed technique involves stimulating the cutaneous receptive field formed by the auricular branch of the vagus nerve in the outer ear, with resulting activation of vagal connections to central and peripheral nervous systems. Increasing evidence indicates that maladaptive neural plasticity may underlie the pathology of several pediatric neurodevelopmental and psychiatric disorders, such as autism spectrum disorder, attention deficit hyperactivity disorder, disruptive behavioral disorder and stress-related disorder. Vagal stimulation may therefore provide a useful intervention for treating maladaptive neural plasticity. In the current review we summarize the current literature primarily on therapeutic use in adults and discuss the prospects of applying taVNS as a therapeutic intervention in specific pediatric neurodevelopmental and other psychiatric disorders. Furthermore, we also briefly discuss factors that would help optimize taVNS protocols in future clinical applications. We conclude from these initial findings that taVNS may be a promising alternative treatment for pediatric disorders which do not respond to other interventions.

The study of moral judgements often centres on moral dilemmas in which options consistent with deontological perspectives (that is, emphasizing rules, individual rights and duties) are in conflict with options consistent with utilitarian judgements (that is, following the greater good based on consequences). Greene et al. (2009) showed that psychological and situational factors (for example, the intent of the agent or the presence of physical contact between the agent and the victim) can play an important role in moral dilemma judgements (for example, the trolley problem). Our knowledge is limited concerning both the universality of these effects outside the United States and the impact of culture on the situational and psychological factors affecting moral judgements. Thus, we empirically tested the universality of the effects of intent and personal force on moral dilemma judgements by replicating the experiments of Greene et al. in 45 countries from all inhabited continents. We found that personal force and its interaction with intention exert influence on moral judgements in the US and Western cultural clusters, replicating and expanding the original findings. Moreover, the personal force effect was present in all cultural clusters, suggesting it is culturally universal. The evidence for the cultural universality of the interaction effect was inconclusive in the Eastern and Southern cultural clusters (depending on exclusion criteria). We found no strong association between collectivism/individualism and moral dilemma judgements. Including participants from 45 countries, Bago et al. find that the situational factors that affect moral reasoning are shared across countries, with diminished observed cultural variation., Comunidad de Madrid [2016-T1/SOC-1395, 2020-5A/SOC-19723]; AEI [PSI2017-85159-P]; UE/FEDER; National Science Centre, Poland [2017/01/X/HS6/01332, 2015/19/D/HS6/00641, 2019/35/B/HS6/00528]; Aarhus University Research Foundation (AUFF) [AUFF-E-2019-9-4]; ANR [ANR-17-EURE-0010]; ANR Labex IAST; Australian Research Council [DP180102384]; CAPES [88887.364180/2019-00]; Carlsberg Foundation [CF16-0444]; Independent Research Fund Denmark [7024-00057B]; Germany's Excellence Strategy [EXC 2126/1-390838866]; FONDECYT, CONICYT [11190980]; German Research Foundation [FOR-2150, LA 3566/1-2]; JSPS [18K03010]; JSPS KAKENHI [16H03079, 17H00875, 18K12015, 20H04581, 17H06342, 20K03479, 20KK0054, 20J21976]; Natural Science Foundation of China (NSFC) [31530032]; Key Technological Projects of Guangdong Province [2018B030335001]; National Science Foundation Graduate Research Fellowship [DGE-1839285]; Polish National Science Center; DFG Beethoven [2016/23/G/HS6/01775]; Portuguese National Foundation for Science and Technology [SFRH/BD/126304/2016, UID/PSI/03125/2019]; PRIN 2017 (Italian Ministry of Education and Research) [20178293XT]; PSA 006 BRA 008 Data Collection in Support of PSADM 001 Measurement Invariance Project; Foundation for Polish Science (START); National Science Centre [2020/36/T/HS6/00256, 2019/33/N/HS6/00054]; Slovak Research and Development Agency [APVV-18-0140, APVV-17-0418, PRIMUS/20/HUM/009, APVV-17-0596]; Social Sciences and Humanities Research Council of Canada [950-224884]; Swedish Research Council [2016-06793]; Project of Philosophy and Social Science Research in Colleges and Universities in Jiangsu Province [2020SJA0017], M.A. Vadillo was supported by 2016-T1/SOC-1395 and 2020-5A/SOC-19723 from Comunidad de Madrid, PSI2017-85159-P from AEI and UE/FEDER. M.P.-C. was supported by 2017/01/X/HS6/01332 from the National Science Centre, Poland. P.M. was supported by Aarhus University Research Foundation (AUFF), starting grant: AUFF-E-2019-9-4. B. Bago was supported by ANR grant ANR-17-EURE-0010 (Investissements d'Avenir programme) and ANR Labex IAST. R.M.R. was supported by the Australian Research Council (DP180102384). N.B.D. was supported by CAPES grant no. 88887.364180/2019-00. C.S., K.A.. and I. Zettler were supported by the Carlsberg Foundation (CF16-0444) and the Independent Research Fund Denmark (7024-00057B). J.L. was supported by EXC 2126/1-390838866 under Germany's Excellence Strategy. K.B. was supported by the following grants from the National Science Centre, Poland: (1) while working on the data collection, no. 2015/19/D/HS6/00641, (2) while working on the final version of the paper, no. 2019/35/B/HS6/00528. A.W. was supported by FONDECYT 11190980, CONICYT. A. Fleischmann was supported by the German Research Foundation (research unit grant FOR-2150, LA 3566/1-2). H.Y. was supported by JSPS grant 18K03010. Y.Y. was supported by JSPS KAKENHI (16H03079, 17H00875, 18K12015 and 20H04581). K.Q. was supported by JSPS KAKENHI (17H06342, 20K03479 and 20KK0054). A. Ikeda was supported by JSPS KAKENHI (20J21976). K.M.K. was supported by National Natural Science Foundation of China (NSFC) grant no. 31530032 and Key Technological Projects of Guangdong Province grant no. 2018B030335001. J.B.C. was supported by National Science Foundation Graduate Research Fellowship grant no. DGE-1839285. M. Parzuchowski, K. Rybus and N.M.S. were supported by Polish National Science Center and DFG Beethoven grant 2016/23/G/HS6/01775. A.C.S. was supported by Portuguese National Foundation for Science and Technology grant no. SFRH/BD/126304/2016. L. Boncinelli was supported by PRIN 2017 grant no. 20178293XT (Italian Ministry of Education and Research). M.F.F.R. was supported by PSA 006 BRA 008 Data Collection in Support of PSADM 001 Measurement Invariance Project. M. Misiak was supported by a scholarship from the Foundation for Polish Science (START) and by a scholarship from the National Science Centre (2020/36/T/HS6/00256). P.B. was supported by Slovak Research and Development Agency project no. APVV-18-0140. M.A. was supported by Slovak Research and Development Agency project no. APVV-17-0418 and project PRIMUS/20/HUM/009. A. Findor and M.H. were supported by the Slovak Research and Development Agency under contract no. APVV-17-0596. T.G. was supported by the Social Sciences and Humanities Research Council of Canada (no. 950-224884). P.P. was supported by the Swedish Research Council (2016-06793). Y.L. was supported by The Project of Philosophy and Social Science Research in Colleges and Universities in Jiangsu Province (grant no. 2020SJA0017). M. Kowal was supported by a scholarship from the National Science Centre (2019/33/N/HS6/00054). P.A. was supported by UID/PSI/03125/2019 from the Portuguese National Foundation for Science and Technology. The funders had no role in study design, data collection and analysis, decision to publish or preparation of the manuscript.