Research into the brain basis of maternal care in mammals identified a set of subcortical limbic structures, which, primed by the oxytocin surge during labor, triggers the species-typical caregiving behaviors that usher young to social living (Feldman, 2015a; Numan and Young, 2016). These brief moments of social contact when mothers express the prototypical behavior of their species carry profound effect on infant sociality; reorganize the infant's lifetime oxytocin system (Champagne et al., 2001; Feldman, 2016; Francis, 1999; Krol et al., 2019), augment the salience of social cues (Marlin et al., 2015), and sculpt the infant's brain and behavior to life within the social ecology (Hammock, 2015). While few functions are as conserved as maternal care, in humans the subcortical structures that underpin mammalian mothering expanded to include insulo-cingulate, temporal, and frontal regions which coalesce to form the ‘human caregiving network’ (Feldman, 2017). Activation of this network supports the human-specific caregiving behavior and enables parents to perform the complex task of preparing human children to life within cultural communities; empathize with the infant's emotion, mentalize to infer infant intentions, prioritize caregiving activities, and plan for long-term parenting goals based on culturally-transmitted social values (Feldman, 2015a; Feldman, 2017).

Consolidation of the human caregiving network during the postpartum months is impacted by the mother's physiological and mental state and links with circulating oxytocin (OT) (Atzil et al., 2011) and cortisol (Seth et al., 2016) levels, degree of maternal anxiety and depression (Pawluski et al., 2017), and representations of own caregiving (Kim et al., 2010). Activation of the caregiving network also underpins the expression of mother-infant behavioral synchrony, the temporally matched repetitive-rhythmic social play which is observed universally and marked by episodes of shared gaze, mutual positive affect, and ‘motherese’ high-pitched vocalizations (Feldman, 2007). Albeit brief, these precious social moments of synchrony expose mothers and infants to massive amounts of social inputs, require coordinated behavior to regulate the high positive arousal, and carry profound effects on infant sociality (Tronick, 1989). Longitudinal studies have shown that mother-infant synchrony plays an important role in children's socialization, emotion regulation, and stress management (Feldman et al., 2010b; Feldman et al., 2013). Furthermore, organization of the parent's caregiving network in infancy shapes children's social-emotional skills as mediated by behavioral synchrony and parental OT (Abraham et al., 2018; Abraham et al., 2016; Kim et al., 2015), highlighting the links between the three components of bonding; the caregiving network, OT system, and synchrony. Indeed, when bonding is disrupted, due to conditions such as postpartum depression or environmental stress, deficits are observed in all three components; activation of the parent's caregiving network, synchronous parenting, and OT production and these carry long-term effects on the child's propensity for psychopathology and maturation of neural systems that underpin empathy and attachment (Davis et al., 2017; Kim et al., 2016; Levy et al., 2019; Pratt et al., 2019).

Across species, the ‘maternal care’ envelope marks the overall provisions transmitted from one generation to the next that contain the evolutionary-acquired information necessary for survival and program the infant's brain to what it means to be a member of that species (Kundakovic and Champagne, 2015; Meaney, 2001). Maternal care comprises a range of long and arduous activities, such as nest building, food retrieval, and, in some primate species, group collaborative and defensive activities (Hayes, 2000; Russell, 2003). Episodes of maternal social contact interfacing with an individual infant are brief, and, in some species, last no longer than several minutes per day for several days (González-Mariscal, 2007; Lucion and Bortolini, 2014). In humans, moments of direct maternal-infant social contact are similarly brief and occupy a fraction of the overall maternal caregiving. Between three and nine months, the sensitive period for social development, episodes of mother-infant face-to-face synchrony typically last 3–5 min, but their impact is long-lasting (Cohn and Tronick, 1988; Feldman, 2015b). One mechanism that underpins the long-term effects of these brief social moments is bio-behavioral synchrony (Feldman, 2017). Moments of mother-infant behavioral synchrony provide a template for the coordination of physiological processes, allowing the mature brain to externally regulate the infant's brain and tune it to social living (Hofer, 1994; Leong et al., 2017). During synchronous play, mothers and infants coordinate their heart rhythms (Feldman et al., 2011b), oxytocin response (Feldman et al., 2010a), and neural oscillations (Leong et al., 2019), and these carry an ‘imprinting-like’ effect on the infant's brain. It is thus likely that these intense social moments also have a distinct signature in the maternal network.

In the current study, we examined whether mother-infant social moments marked by increased synchrony would trigger increased activations across the caregiving network in postpartum mothers. The human caregiving network comprises the inter-connected functioning of its subcortical (amygdala, VTA), para-limbic (AI, ACC), temporal (STS/STG, TP), and frontal (mPFC) components into a functional network that coalesces to support human caregiving (Feldman, 2015a; Feldman et al., 2019; Kim et al., 2016; Swain et al., 2014). While the brain of any adult exhibits responses in regions of this network to infant cues (Kringelbach et al., 2008; Rilling and Mascaro, 2017), synchronous social moments are expected to activate a coherent response across the network that is stronger and more unified as compared to similar maternal-infant cues that do not contain a social component. Although such comparison has not yet been tested, it is reasonable to assume that since maternal care is a time-consuming, metabolically-costly endeavor, bearing critical impact on species continuity, the mother's brain would not activate to it full capacity when resources are needed for other tasks but would cohere to its full expression to sustain these brief moments of sociality.

In addition, we examined whether activations of the human caregiving network to social moments would show greater sensitivity to oxytocin administration as compared to similar episodes of maternal-infant presence that do not contain a social component. OT is an important modulator of the brain's social functions (Zink and Meyer-Lindenberg, 2012) and supports reorganization of the mother's brain following childbirth (Insel and Young, 2001). OT plays a critical role in neural plasticity at the molecular and network assembly levels and such plasticity augments the salience and reward value of the infant to its mother (Marlin et al., 2015; Oettl et al., 2016; Valtcheva and Froemke, 2019), and both experimental and knockout studies demonstrate the causal role of OT in the initiation of maternal social behavior (Higashida et al., 2010; Lopatina et al., 2012). Human studies have shown that peripheral OT levels are associated with mother-infant synchrony (Feldman et al., 2011a) and activation of the human caregiving network (Abraham et al., 2014; Rilling and Mascaro, 2017). Intranasal OT administration affects parenting by increasing parents' prototypical-rhythmic behaviors and augmenting parent-child synchrony (Naber et al., 2010; Weisman et al., 2012a) and OT may thus target the social context which is marked by high synchrony.

While we expected OT administration to affect primarily neural response to the social condition, the direction of its effects remained a research question. The effects of OT administration on BOLD response are far from clear and the literature is mixed on whether OT increases or decreases activations of nodes within the caregiving network (Chen et al., 2017; Grace et al., 2018; Martins et al., 2020; Wang et al., 2017; Wigton et al., 2015). Conceptually, whereas the ‘social salience’ hypothesis argues that OT increases the salience of social stimuli and to enhanced response to social signals (Shamay-Tsoory and Abu-Akel, 2016), the anxiolytic model on OT (Neumann and Slattery, 2016) may suggest that OT would level-out the increased response to social-emotional cues to maintain equilibrium and calm. Of the few studies that examined the effects of OT on the parent's neural response to infant stimuli, several indicated attenuation of BOLD response under OT. Wittfoth-Schardt et al., 2012 tested fathers' neural response to unfamiliar, familiar, and own infant pictures and found increased response in subcortical reward regions, hippocampus, AI, STS, and OFC to own infant under placebo (PBO), which attenuated under OT, and concluded that OT attenuates neural response as a function of social salience. Bos et al., 2018, testing mothers, similarly showed increased response to own infant pictures under PBO, which decreased under OT in VTA, putamen, and amygdala and concluded that OT attenuates neural response as a function of social arousal. Riem et al., 2016 showed amygdala attenuation under OT to infant cries pending maternal attachment representations. These studies lend support to the hypothesis that while the social condition would increase activations in the caregiving network under PBO, OT may level out these socially driven activations marked by salience and arousal. However, since other studies showed BOLD increases under OT in fathers' brain (Li et al., 2017) and as the current consensus is that OT effects are time-, person-, and context-sensitive (Bartz et al., 2011), we hypothesize that OT would target the social condition and explored the direction of its effects.

To describe mothers' neural responses to synchronous social moments, we expanded on a well-researched paradigm into the parental brain that utilized presentation of individually-tailored stimuli collected in the home ecology (Atzil et al., 2011; Elmadih et al., 2016; Noriuchi et al., 2008). We included three separate conditions that depict typical mother-child social and non-social contexts in the home environment. Across conditions, mothers were filmed sitting next to their child in the same level and distance, to control for differences in physical proximity and posture. In the first condition, mothers sat next to their infant while being otherwise engaged (Condition I, Unavailable); in the second, mothers sat facing the infant but did not engage in social interactions (Condition II, Unresponsive); in the third, mothers engaged in a prototypical, rhythmic social play of peek-a-boo (Condition III: Social). Mothers were imaged twice in a double-blind within-subject placebo-control design and observed the same three conditions viewing themselves (Self) and an unfamiliar mother-infant dyad (Other) once following administration of oxytocin (OT) and once after placebo (PBO).

We expected that the social condition would elicit greater response as compared to the other conditions across the caregiving network. Similarly, we hypothesized that OT would impact specifically the caregiving network's response to the social condition, and tested whether these OT-mediated responses would follow the ‘social salience’ hypothesis (i.e., increased brain activations to social condition under OT) or the anxiolytic model of OT (decreased brain activation under OT). In addition, and as an open research question, we explored whether activation of the caregiving network to the social condition would show greater consistency between the two imaging sessions as compared to the other conditions, particularly in the insula and structures of the temporal cortex (STS, TP). The social condition is characterized by repetitive-rhythmic social stimuli that may trigger distinct activations in the ‘sociotemporal brain’ (Schirmer et al., 2016), which gauges durations, patterns, and frequencies of temporally-ordered stimuli. We thus investigated whether the synchronous moments of mother-infant social play would elicit greater temporal consistency in limbic, insular, and temporal regions that underpin the brain's perception of temporal regularities. Finally, we expected that the mother's neural activations would show a differential response between her own and an unfamiliar infant and that these would increase in the social condition. To investigate if our results are specific to the caregiving network, we examined the effects of the social condition on two additional networks; the default mode network (DMN), a well-described network known to be activated by self-related processing (Buckner et al., 2008; Peer et al., 2015; Salomon et al., 2014; Spreng et al., 2009), and the visual network, an occipital task-positive network unrelated to bonding. We expected that while the DMN may show self-related effects, both the DMN and the visual neural systems would not be sensitive to the social versus non-social conditions or to OT administration, highlighting the specific response of the human caregiving network to social cues.

For social mammals born with an immature brain that requires the external regulation of the mother for growth and development, moments of maternal-infant social interactions hold a special significance. These brief social moments integrate multiple well-orchestrated bio-behavioral processes that consolidate the caregiving network in the maternal brain, trigger the species-typical caregiving behavior, and carry long-term impact on the developing infant brain (Feldman, 2020). Our study uniquely tests the response of the human caregiving network in the maternal brain to these social moments, versus other moments of non-social mother-infant presence, to shed further light on how human mothers' brains may change following birth and vary with the caregiving experience with one's infant.

To tap the response of the caregiving network to social interactions, we employed a double-blind, within subject, OT/placebo crossover design; hence, one of the novel aspects of our study is the closely-scheduled repeated imaging of the mother's brain in response to multiple ecological contexts during the sensitive period of bond formation. Findings describe several processes by which social moments impact the maternal network. First, whole-brain analysis (Figures 2 and 3) demonstrated a significant widespread response to the social condition across large expanses of the brain, including the human caregiving network. This substantial and integrated response across temporal, frontal, and insular cortices, as well as subcortical regions, was specific to the social context, which elicited much larger neural activations in comparison with both the ‘unavailable’ and ‘unresponsive’ conditions. While these conditions included familiar mother-infant daily stimuli, a similar degree of physical proximity, and the same mother-infant sitting posture, they did not include the synchronous social component (see Figure 1). Moreover, the specific activations to the social condition was not found for the other systems we tested: the DMN, which underpins self-related processes, and the low level visual system regions, suggesting that our findings do not merely reflect a widespread neural response but are specific to the human caregiving network. These findings support our first hypothesis that these brief and universal synchronous social moments uniquely elicit a widespread response in the caregiving network, which sustain human maternal care, and that such differentiation of social from non-social mother-child episodes is specific to this network.

Second, we show that the effects of OT on the caregiving network were more notable during the social condition. OT decreased brain activation to the social condition throughout the caregiving network, indicating that the mother's neural response to synchronous social moments is more sensitive to OT. Bayesian analysis provided support for a coherent network-wide response to the social condition under OT across the caregiving network regions of interest (see Figure 4), further supporting the proposition that the interaction of social cues and OT impacts the network-wide functioning of this network. The special sensitivity of these socially driven neural activations to OT administration showed a consistent pattern which was similar across the pre-registered ROIs comprising the caregiving network (see Figure 4B). While these regions increased activity to the social context under natural conditions (i.e. PBO), OT leveled out these socially driven activations and under OT no significant differences were found between the social and non-social conditions.

To date, studies on the effects of OT administration on BOLD response have yielded mixed results and no straightforward explanation. Still, our findings are consistent with the majority of prior research which showed that OT targets social functions. OT enhances social behaviors, increases social collaboration, and augments understanding of social cues, and OT's effects on the brain are sensitive to social salience (Di Simplicio et al., 2009; Guastella et al., 2008; Hurlemann et al., 2010). Our findings are also consistent with research on OT's effects on the parental brain, which showed increased activation to own-infant pictures under PBO across wide areas of the caregiving network that were attenuated under OT, and the authors considered these results to stem from the social salience and high arousal embedded in these stimuli (Bos et al., 2018; Wittfoth-Schardt et al., 2012). A recent study testing the effects of OT administration via both intravenous and intranasal pathways on neural response during resting state showed that OT via both pathways decreased activity in amygdala, insula, TP, and parahippocampal gyrus, and the decrease was mediated by elevations in peripheral OT levels (Martins et al., 2020). Our findings similarly show a combination of increase in peripheral OT levels following OT administration and attenuation of neural response to social cues in the same regions. Our results may extend the resting state findings to include the caregiving network's processing of ongoing live social stimuli. In combination, these results may suggest that one pathway for the anxiolytic effects of OT may relate to the attenuation of activity in a limbic network that monitors salience cues, gauges danger signals, and integrates exteroceptive and interoceptive inputs to give immediacy and focus to ongoing events.

While our findings may be consistent with some prior research, it is important to note that OT's effects on BOLD response is complex and there is probably no simple relationship between social significance and consistent increase or decrease in levels of activations. To date, the effects of OT administration on the human brain have mainly been tested via alterations in BOLD response; however, much further research is needed along several additional dimensions and the application of more finely-tuned analytic tools. OT effects may, for instance, express in augmenting the brain's sensitivity to temporal regularities during the processing of social stimuli and further research should examine the conditions, specific patterns, and populations under which such sensitivity is enhanced or suppressed.

One possible cause for the associations between OT and the processing of temporal patterns in the mother's brain may relate to the crosstalk between OT and dopamine (DA) that underpins bonding. The initiation of maternal care in rodents involves a two-step process; first, OT leads to long-term depression in amygdala to suppress social avoidance of infant stimuli (Gur et al., 2014), and next, OT connects with DA through striatal neurons that encode for both OT and dopamine D1 receptors (Olazábal and Young, 2006). This enables DA neurons that encode sensory-motor general reward patterns to also encode the temporal patterns of social reward (Báez-Mendoza and Schultz, 2013; Ross and Young, 2009). This process allows the brain to internalize the social partner and its preferences, encode relationship-specific socio-temporal patterns, and draw reward from the matching of the social actions of self and partner, that is, social synchrony (Báez-Mendoza and Schultz, 2013; Schultz, 2016). The crosstalk of OT and DA triggered by maternal care enables the mother's brain to integrate rewarding experiences from the infant's smell, touch, babbling, and cute face into an overall representation that contains the dyad-specific temporal rhythms to sustain the attachment bond (Ross et al., 2009).

In addition to reward, OT also has well-known anxiolytic properties (Neumann, 2008) and in the context of maternal care the soothing function of OT may be important for survival. During labor, OT surges to levels much higher than the body's daily levels and these function to sooth the mother's pain and stress through the regulatory effects of OT on the hypothalamic-pituitary-adrenal-axis and sympathetic arousal (Carter, 2014; Neumann and Slattery, 2016). OT enables tranquility during the birth process, attenuates amygdala response to external events, and diminishes insular monitoring of unfamiliar interoceptive signals, keeping mother's brain from oscillating between extreme emotional states. Since OT administration leads to unusually high peripheral OT levels (Weisman et al., 2012b), the anxiolytic properties of OT during the postpartum period, particularly in response to infant stimuli, may have functioned here in a similar way to level-out the mother's neural response to different emotional states. However, this hypothesis is preliminary and requires much further research, possibly by using stimuli that target mothering-related stress and anxiety.

Our exploratory WSC analysis showed greater consistency in patterns of activation across the two imaging sessions in the social condition. Such socially driven consistency was found across the entire maternal caregiving network and a Bayesian analysis supported a network-wide temporal consistency across ROIs, particularly in amygdala, insula, and TP. It is important to emphasize that the WSC analyses were exploratory and should be interpreted cautiously. A repeated and highly arousing stimulus may in of itself elicit greater consistency across imaging sessions as compared to the other conditions that were more boring, less salient, and may generally entrain neural processes to a lesser degree. Similarly, the temporal consistency did not show differences related to own versus unfamiliar interaction and we did not tease apart conditions of infant alone, mother alone, and maternal-infant presence. Hence, these findings should be considered as very preliminary and future research should carefully tease apart the components related to salience from those related to self and those related to the mother's own attachment to her infant. Despite their very preliminary nature, the WSC analysis may point several important directions for future research on the neural basis of maternal-infant attachment and the effects of OT administration on the brain more generally. For instance, it appears that while the magnitude of BOLD responses in the caregiving network to social stimuli decreased under OT, the WSC analysis, which encodes the temporal pattern of response, was preserved. This suggests that regional BOLD fluctuations alone may not fully capture the neural processes induced by OT administration during social moments, and future studies may apply more novel multivariate spatial and temporal patterns analyses (Saarimäki et al., 2016; Ulmer Yaniv et al., 2020). Possibly, OT has a complex, multi-dimensional effect on the neural basis of social processes that impacts both the signal magnitude and the encoding of temporal patterns during the processing of social cues and both dimensions should be considered in future studies.

Another interesting outcome of the WSC analysis is pinpointing the specific regions that exhibited the greatest consistency in response to social stimuli; the amygdala, insula, and TP, nodes that have been implicated in the perception of temporal regularities, particularly in social contexts. For instance, amygdala neurons in the primate brain were found to simulate and foreshadow the partner's decision making process (Grabenhorst et al., 2019). Similarly, the insula and regions of the temporal cortex, including the STS and TP, monitor the salience and valance of stimuli and underpin the perception of temporal regularities and the duration and patterning of social stimuli (Schirmer et al., 2016). The insula plays a key role in interoception, the process by which the brain uses repeated experiences of own bodily signals to build predictions of self and others' physical and mental states (Barrett and Simmons, 2015; Salomon et al., 2016; Seth, 2013). Insular activations in the maternal brain enable mothers to represent the infant's bodily signals of hunger, fatigue, satiety, and pain in their own brain (Abraham et al., 2019). The insula also serves as a center of allostasis, the brain resource-regulator function that sustains the mother's ability to detect patterned regularities in the infant's physiological needs and satisfy them in a timely manner before they arise (Schulkin and Sterling, 2019). Much further research is needed to explore the role of temporal regularities in the mothers' limbic, insular, and temporal response to social rhythms and their long-term effects on the infant's developing brain, social behavior, and the formation of the parent-infant attachment.

Although preliminary, we found evidence for brain-behavior coupling between patterns of mother-infant synchrony observed in the home environment and maternal neural response to her infant in the social condition in ACC, insula, and VTA, but not in other conditions. While these findings should be treated with great caution, due to the small sample size, they again show the selectivity of the social condition in linking maternal neural response to the degree of synchrony the infant experiences during daily playful moments in the home environment. Since mother-infant synchrony is an important dyadic experience that is both individually stable from infancy to adulthood and predicts social-emotional competencies, our preliminary findings suggest that this important dyad-specific pattern of synchrony may also be linked with activation of the mother's caregiving network. Such brain-behavior link between behavioral synchrony and mother's neural response to own infant in the social condition appears to be specific to the synchrony dimension of mother-infant interaction. The degree of infant social engagement in the interaction, while showing correlations with maternal VTA and TP response in the social condition, these correlations were not specific to own infant but emerged for both own and unfamiliar social interaction. Notably, the infant's temperamental dimensions of reactivity and emotionality were unrelated to maternal neural response in any condition, corroborating developmental perspectives which emphasize that attachment and temperament are two distinct processes (Bowlby, 1969; Sroufe, 1985). However, since we did not have standard measures to assess temperament, this should be considered a study limitation and our results underscore the need for much further research on the bi-directional associations between mother's brain, mother-infant interaction, and maternal and child factors.

Results indicate that nearly all regions of the caregiving network showed sensitivity to one’s own vs. unfamiliar interactions. Contrary to our hypothesis, the directionality of the BOLD response varied across ROIs; higher activation for the ‘Self’ condition in the ACC and insular regions, and higher activity to the ‘Other’ condition in the other ROIs, with no differences in the VTA. These findings suggest that the caregiving network is highly responsive to the Self-Other distinction and exhibits a differential response to self-related mother-infant stimuli across six of seven ROIs. The ACC and insular regions respond to self-related stimuli across different types of stimuli (Karnath and Baier, 2010; Northoff et al., 2006; Qin et al., 2012; Salomon et al., 2018). Yet, in contrast to our expectation, no interaction effect was found between maternal condition and own-unfamiliar infant, indicating that the mother's neural response to social moments did not differentiate the two. Possibly, the salience of these arousing species-typical social exchanges is especially high and may trump differentiation of self and other, but further research is needed to understand these findings.

Response of the DMN showed a different pattern of activations from the one found for regions of the caregiving network. We included the DMN in our preregistered ROIs to assess the caregiving network in comparison with another well-characterized network known to be sensitive to self-related processing (Andrews-Hanna et al., 2014; Northoff et al., 2006; Peer et al., 2015) and to pinpoint the effects of social moments on this network. The DMN provides a useful comparison as it is thought to sustain the sense of self and is sensitive to self-other distinction across numerous types of stimuli (Davey et al., 2016; Salomon et al., 2009). As expected, the DMN showed a significant effect of Self vs. Other; however, contrary to previous studies (Andrews-Hanna et al., 2014; Northoff et al., 2006; Salomon et al., 2014), it showed higher BOLD responses to the Other condition. This is likely due to the mother's orientation towards external stimuli and possibly as a result of averaging the different regions of the DMN, which have different functional selectivity to self-related stimuli (Araujo et al., 2015; Davey et al., 2016; Northoff et al., 2006; Salomon et al., 2014). Still, while the caregiving network showed decreased activation under OT to social stimuli, no such effects emerged for the DMN. In addition, the DMN did not display consistent temporal patterns in the WSC. Additional analysis of the visual network, a task positive network that was activated by the visual stimulation, similarly showed no differential response for social stimuli under OT, highlighting the specificity of our findings to the maternal caregiving network and to mother-infant social moments.

Limitations of the study include the relatively small number of participants and the relatively high attrition rate, which partly relate to the fact that mothers were imaged twice within a 2-week span during the postpartum and if mothers could not schedule the next meeting or one scan had technical problems the participant was excluded from the study. Similarly, inclusion of more controlled stimuli alongside the ecologically valid ones could have shed further light on the mother's caregiving network. Despite these limitations, our study introduces a novel ecologically-valid paradigm, examines pre-registered ROIs, and integrates multi-measure methodologies, including brain imaging, social behavioral coding, OT administration, and hormonal analysis to expand knowledge on the mother's neural response to social moments.

Characterization of the human mother's brain requires much further study. Whereas our study examined the neural responses of postpartum mothers to social and non-social caregiving experiences, future research may target other caregivers, such as fathers, grandparents, or childcare providers. Similarly, we imaged mothers who are raising infants in typical mother-father families and other family constellations, such as single or same-sex parents require further research.

Future work could rigorously test the cross-generational transmission of human sociality through longitudinal studies that examine linkage between a mother's neural response to social moments with her infant, versus non-social caregiving experiences, and her child's future social-emotional development, psychological well-being, or indices of brain maturation and neural response to social and affiliative cues. Another window into the cross generational transmission is by targeting high-risk conditions known to impact children's social competencies. Neural activations in the maternal brain to infant stimuli are attenuated in conditions such as poverty (Kim et al., 2017) or depression (Kim et al., 2016) and longitudinal studies show that disruptions in maternal synchronous caregiving predict later insensitivity of the child's brain to attachment cues (Pratt et al., 2019). However, a longitudinal study linking a mother's neural response in the postpartum with her child's later brain activations to social stimuli has not been conducted and such research may shed further light on how the mother's neural response to moments of social synchrony plays a particularly important role in tuning the child's brain to social cues. Finally, our findings can inform the construction of specific interventions that target maternal neural response to social interactions and aim to boost the salience and reward of the infant and the attachment relationship. Much further research is needed to fully understand the mechanisms that sustain consolidation of the caregiving network in health and pathology and describe how cultural practices, personal habits, and meaning systems shape the mother's neural response and are then transferred to the infant to cement the transmission of human cultural heritage.

Raw, subject by subject, anonymized brain data (fMRI); group level data (e.g. unthresholded group maps on MNI template) and raw subject by subject data from the ROI analysis (csv and JASP files) are freely available. These files are uploaded to our OSF account (https://osf.io/mszqj/?view_only=0daf10c02c984ead8929452edf44e550) to allow full transparency of the data.

1. 1. Shimon-Raz O
   2. Salomon R
   3. Bloch M
   4. Romano GA
   5. Yeshurun Y
   6. Ulmer-Yaniv A
   7. Zagoory-Sharon O
   8. Feldman R

   (2020) Open Science Framework

   ID mszqj. Mother Brain is Wired for Social Moments.

   https://osf.io/mszqj/?view_only=0daf10c02c984ead8929452edf44e550

1. 1. Abraham E
   2. Hendler T
   3. Shapira-Lichter I
   4. Kanat-Maymon Y
   5. Zagoory-Sharon O
   6. Feldman R

   (2014) Father's brain is sensitive to childcare experiences

   PNAS 111:9792–9797.

   https://doi.org/10.1073/pnas.1402569111

   • PubMed
   • Google Scholar
2. 1. Abraham E
   2. Hendler T
   3. Zagoory-Sharon O
   4. Feldman R

   (2016) Network integrity of the parental brain in infancy supports the development of children's social competencies

   Social Cognitive and Affective Neuroscience 11:1707–1718.

   https://doi.org/10.1093/scan/nsw090

   • PubMed
   • Google Scholar
3. 1. Abraham E
   2. Raz G
   3. Zagoory-Sharon O
   4. Feldman R

   (2018) Empathy networks in the parental brain and their long-term effects on children's stress reactivity and behavior adaptation

   Neuropsychologia 116:75–85.

   https://doi.org/10.1016/j.neuropsychologia.2017.04.015

   • PubMed
   • Google Scholar
4. 1. Abraham E
   2. Hendler T
   3. Zagoory-Sharon O
   4. Feldman R

   (2019) Interoception sensitivity in the parental brain during the first months of parenting modulates children's somatic symptoms six years later: The role of oxytocin

   International Journal of Psychophysiology 136:39–48.

   https://doi.org/10.1016/j.ijpsycho.2018.02.001

   • PubMed
   • Google Scholar
5. 1. Andrews-Hanna JR
   2. Smallwood J
   3. Spreng RN

   (2014) The default network and self-generated thought: component processes, dynamic control, and clinical relevance

   Annals of the New York Academy of Sciences 1316:29–52.

   https://doi.org/10.1111/nyas.12360

   • PubMed
   • Google Scholar
6. 1. Araujo HF
   2. Kaplan J
   3. Damasio H
   4. Damasio A

   (2015) Neural correlates of different self domains

   Brain and Behavior 5:e00409.

   https://doi.org/10.1002/brb3.409

   • PubMed
   • Google Scholar
7. 1. Atzil S
   2. Hendler T
   3. Feldman R

   (2011) Specifying the neurobiological basis of human attachment: brain, hormones, and behavior in synchronous and intrusive mothers

   Neuropsychopharmacology 36:2603–2615.

   https://doi.org/10.1038/npp.2011.172

   • PubMed
   • Google Scholar
8. 1. Báez-Mendoza R
   2. Schultz W

   (2013) The role of the striatum in social behavior

   Frontiers in Neuroscience 7:233.

   https://doi.org/10.3389/fnins.2013.00233

   • PubMed
   • Google Scholar
9. 1. Barrett LF
   2. Simmons WK

   (2015) Interoceptive predictions in the brain

   Nature Reviews Neuroscience 16:419–429.

   https://doi.org/10.1038/nrn3950

   • PubMed
   • Google Scholar
10. 1. Bartz JA
    2. Zaki J
    3. Bolger N
    4. Ochsner KN

    (2011) Social effects of oxytocin in humans: context and person matter

    Trends in Cognitive Sciences 15:301–309.

    https://doi.org/10.1016/j.tics.2011.05.002

    • PubMed
    • Google Scholar
11. 1. Benjamini Y
    2. Hochberg Y
    3. Benjamini YHY

    (1995) Benjamini and Y fdr.pdf

    J R Stat Soc Ser B 57:289–300.

    https://doi.org/10.2307/2346101

    • Google Scholar
12. 1. Bos PA
    2. Spencer H
    3. Montoya ER

    (2018) Oxytocin reduces neural activation in response to infant faces in nulliparous young women

    Social Cognitive and Affective Neuroscience 13:1099–1109.

    https://doi.org/10.1093/scan/nsy080

    • Google Scholar
13. Book

    1. Bowlby J

    (1969)

    Attachment and Loss

    Basic Books.

    • Google Scholar
14. 1. Buckner RL
    2. Andrews-Hanna JR
    3. Schacter DL

    (2008) The brain's default network: anatomy, function, and relevance to disease

    Annals of the New York Academy of Sciences 1124:.

    https://doi.org/10.1196/annals.1440.011

    • PubMed
    • Google Scholar
15. 1. Carter CS

    (2014) Oxytocin pathways and the evolution of human behavior

    Annual Review of Psychology 65:17–39.

    https://doi.org/10.1146/annurev-psych-010213-115110

    • PubMed
    • Google Scholar
16. 1. Champagne F
    2. Diorio J
    3. Sharma S
    4. Meaney MJ

    (2001) Naturally occurring variations in maternal behavior in the rat are associated with differences in estrogen-inducible central oxytocin receptors

    PNAS 98:12736–12741.

    https://doi.org/10.1073/pnas.221224598

    • PubMed
    • Google Scholar
17. 1. Chen X
    2. Gautam P
    3. Haroon E
    4. Rilling JK

    (2017) Within vs. between-subject effects of intranasal oxytocin on the neural response to cooperative and non-cooperative social interactions

    Psychoneuroendocrinology 78:22–30.

    https://doi.org/10.1016/j.psyneuen.2017.01.006

    • PubMed
    • Google Scholar
18. 1. Cohn JF
    2. Tronick EZ

    (1988) Mother-infant face-to-face interaction: influence is bidirectional and unrelated to periodic cycles in either partner's behavior

    Developmental Psychology 24:386–392.

    https://doi.org/10.1037/0012-1649.24.3.386

    • Google Scholar
19. 1. Cox JL
    2. Holden JM
    3. Sagovsky R

    (1987) Detection of postnatal depression development of the 10-item edinburgh postnatal depression scale

    The British Journal of Psychiatry : The Journal of Mental Science 150:782–786.

    https://doi.org/10.1192/bjp.150.6.782

    • PubMed
    • Google Scholar
20. 1. Davey CG
    2. Pujol J
    3. Harrison BJ

    (2016) Mapping the self in the brain's default mode network

    NeuroImage 132:390–397.

    https://doi.org/10.1016/j.neuroimage.2016.02.022

    • PubMed
    • Google Scholar
21. 1. Davis EP
    2. Stout SA
    3. Molet J
    4. Vegetabile B
    5. Glynn LM
    6. Sandman CA
    7. Heins K
    8. Stern H
    9. Baram TZ

    (2017) Exposure to unpredictable maternal sensory signals influences cognitive development across species

    PNAS 114:10390–10395.

    https://doi.org/10.1073/pnas.1703444114

    • PubMed
    • Google Scholar
22. 1. Di Simplicio M
    2. Massey-Chase R
    3. Cowen PJ
    4. Harmer CJ

    (2009) Oxytocin enhances processing of positive versus negative emotional information in healthy male volunteers

    Journal of Psychopharmacology 23:241–248.

    https://doi.org/10.1177/0269881108095705

    • PubMed
    • Google Scholar
23. 1. Elmadih A
    2. Wan MW
    3. Downey D
    4. Elliott R
    5. Swain JE
    6. Abel KM

    (2016) Natural variation in maternal sensitivity is reflected in maternal brain responses to infant stimuli

    Behavioral Neuroscience 130:500–510.

    https://doi.org/10.1037/bne0000161

    • PubMed
    • Google Scholar
24. Conference

    1. Evans AC
    2. Collins DL
    3. Mills SR
    4. Brown ED
    5. Kelly RL
    6. Peters TM

    (1994) 3D statistical neuroanatomical models from 305 MRI volumes

    IEEE Nuclear Science Symposium & Medical Imaging Conference.

    https://doi.org/10.1109/NSSMIC.1993.373602

    • Google Scholar
25. Book

    1. Feldman R

    (1998)

    Coding Interactive Behavior Manual

    Bar-Ilan Univ Press.

    • Google Scholar
26. 1. Feldman R

    (2007) Parent?infant synchrony and the construction of shared timing; physiological precursors, developmental outcomes, and risk conditions

    Journal of Child Psychology and Psychiatry 48:329–354.

    https://doi.org/10.1111/j.1469-7610.2006.01701.x

    • Google Scholar
27. 1. Feldman R
    2. Gordon I
    3. Zagoory-Sharon O

    (2010a) The cross-generation transmission of oxytocin in humans

    Hormones and Behavior 58:669–676.

    https://doi.org/10.1016/j.yhbeh.2010.06.005

    • PubMed
    • Google Scholar
28. 1. Feldman R
    2. Singer M
    3. Zagoory O

    (2010b) Touch attenuates infants' physiological reactivity to stress

    Developmental Science 13:271–278.

    https://doi.org/10.1111/j.1467-7687.2009.00890.x

    • PubMed
    • Google Scholar
29. 1. Feldman R
    2. Gordon I
    3. Zagoory-Sharon O

    (2011a) Maternal and paternal plasma, salivary, and urinary oxytocin and parent-infant synchrony: considering stress and affiliation components of human bonding

    Developmental Science 14:752–761.

    https://doi.org/10.1111/j.1467-7687.2010.01021.x

    • PubMed
    • Google Scholar
30. 1. Feldman R
    2. Magori-Cohen R
    3. Galili G
    4. Singer M
    5. Louzoun Y

    (2011b) Mother and infant coordinate heart rhythms through episodes of interaction synchrony

    Infant Behavior and Development 34:569–577.

    https://doi.org/10.1016/j.infbeh.2011.06.008

    • PubMed
    • Google Scholar
31. Book

    1. Feldman R

    (2012) Parenting Behavior as the Environment Where Children Grow

    Cambridge University Press.

    https://doi.org/10.1017/CBO9781139016827.031

    • Google Scholar
32. 1. Feldman R
    2. Bamberger E
    3. Kanat-Maymon Y

    (2013) Parent-specific reciprocity from infancy to adolescence shapes children's social competence and dialogical skills

    Attachment & Human Development 15:407–423.

    https://doi.org/10.1080/14616734.2013.782650

    • PubMed
    • Google Scholar
33. 1. Feldman R

    (2015a) The adaptive human parental brain: implications for children's social development

    Trends in Neurosciences 38:387–399.

    https://doi.org/10.1016/j.tins.2015.04.004

    • PubMed
    • Google Scholar
34. 1. Feldman R

    (2015b) Sensitive periods in human social development: new insights from research on oxytocin, synchrony, and high-risk parenting

    Development and Psychopathology 27:369–395.

    https://doi.org/10.1017/S0954579415000048

    • PubMed
    • Google Scholar
35. 1. Feldman R

    (2016) The neurobiology of mammalian parenting and the biosocial context of human caregiving

    Hormones and Behavior 77:3–17.

    https://doi.org/10.1016/j.yhbeh.2015.10.001

    • PubMed
    • Google Scholar
36. 1. Feldman R

    (2017) The neurobiology of human attachments

    Trends in Cognitive Sciences 21:80–99.

    https://doi.org/10.1016/j.tics.2016.11.007

    • PubMed
    • Google Scholar
37. 1. Feldman R
    2. Braun K
    3. Champagne FA

    (2019) The neural mechanisms and consequences of paternal caregiving

    Nature Reviews Neuroscience 20:205–224.

    https://doi.org/10.1038/s41583-019-0124-6

    • PubMed
    • Google Scholar
38. 1. Feldman R

    (2020) What is resilience: an affiliative neuroscience approach

    World Psychiatry 19:132–150.

    https://doi.org/10.1002/wps.20729

    • PubMed
    • Google Scholar
39. 1. Feldman R
    2. Eidelman AI

    (2003) Direct and indirect effects of breast milk on the neurobehavioral and cognitive development of premature infants

    Developmental Psychobiology 43:109–119.

    https://doi.org/10.1002/dev.10126

    • PubMed
    • Google Scholar
40. 1. Feldman R
    2. Eidelman AI

    (2007) Maternal postpartum behavior and the emergence of infant-mother and infant-father synchrony in preterm and full-term infants: the role of neonatal vagal tone

    Developmental Psychobiology 49:290–302.

    https://doi.org/10.1002/dev.20220

    • PubMed
    • Google Scholar
41. 1. Francis D

    (1999) Nongenomic transmission across generations of maternal behavior and stress responses in the rat

    Science 286:1155–1158.

    https://doi.org/10.1126/science.286.5442.1155

    • Google Scholar
42. 1. Goebel R
    2. Esposito F
    3. Formisano E

    (2006) Analysis of functional image analysis contest (FIAC) data with brainvoyager QX: from single-subject to cortically aligned group general linear model analysis and self-organizing group independent component analysis

    Human Brain Mapping 27:392–401.

    https://doi.org/10.1002/hbm.20249

    • PubMed
    • Google Scholar
43. Book

    1. Goldsmith HH
    2. Rothbart MK

    (1996)

    The Laboratory Temperament Assessment Battery (LAB-TAB): Prelocomotor Version 3.0

    University of Wisconsin.

    • Google Scholar
44. 1. González-Mariscal G

    (2007) Mother rabbits and their offspring: timing is everything

    Developmental Psychobiology 49:71–76.

    https://doi.org/10.1002/dev.20196

    • PubMed
    • Google Scholar
45. 1. Gordon I
    2. Vander Wyk BC
    3. Bennett RH
    4. Cordeaux C
    5. Lucas MV
    6. Eilbott JA
    7. Zagoory-Sharon O
    8. Leckman JF
    9. Feldman R
    10. Pelphrey KA

    (2013) Oxytocin enhances brain function in children with autism

    PNAS 110:20953–20958.

    https://doi.org/10.1073/pnas.1312857110

    • PubMed
    • Google Scholar
46. 1. Grabenhorst F
    2. Báez-Mendoza R
    3. Genest W
    4. Deco G
    5. Schultz W

    (2019) Primate amygdala neurons simulate decision processes of social partners

    Cell 177:986–998.

    https://doi.org/10.1016/j.cell.2019.02.042

    • PubMed
    • Google Scholar
47. 1. Grace SA
    2. Rossell SL
    3. Heinrichs M
    4. Kordsachia C
    5. Labuschagne I

    (2018) Oxytocin and brain activity in humans: a systematic review and coordinate-based meta-analysis of functional MRI studies

    Psychoneuroendocrinology 96:6–24.

    https://doi.org/10.1016/j.psyneuen.2018.05.031

    • PubMed
    • Google Scholar
48. 1. Granat A
    2. Gadassi R
    3. Gilboa-Schechtman E
    4. Feldman R

    (2017) Maternal depression and anxiety, social synchrony, and infant regulation of negative and positive emotions

    Emotion 17:11–27.

    https://doi.org/10.1037/emo0000204

    • PubMed
    • Google Scholar
49. 1. Guastella AJ
    2. Mitchell PB
    3. Dadds MR

    (2008) Oxytocin increases gaze to the eye region of human faces

    Biological Psychiatry 63:3–5.

    https://doi.org/10.1016/j.biopsych.2007.06.026

    • PubMed
    • Google Scholar
50. 1. Gur R
    2. Tendler A
    3. Wagner S

    (2014) Long-term social recognition memory is mediated by oxytocin-dependent synaptic plasticity in the medial amygdala

    Biological Psychiatry 76:377–386.

    https://doi.org/10.1016/j.biopsych.2014.03.022

    • PubMed
    • Google Scholar
51. 1. Hammock EA

    (2015) Developmental perspectives on oxytocin and vasopressin

    Neuropsychopharmacology 40:24–42.

    https://doi.org/10.1038/npp.2014.120

    • PubMed
    • Google Scholar
52. 1. Hayes LD

    (2000) To nest communally or not to nest communally: a review of rodent communal nesting and nursing

    Animal Behaviour 59:677–688.

    https://doi.org/10.1006/anbe.1999.1390

    • PubMed
    • Google Scholar
53. 1. Higashida H
    2. Lopatina O
    3. Yoshihara T
    4. Pichugina YA
    5. Soumarokov AA
    6. Munesue T
    7. Minabe Y
    8. Kikuchi M
    9. Ono Y
    10. Korshunova N
    11. Salmina AB

    (2010) Oxytocin signal and social behaviour: comparison among adult and infant oxytocin, oxytocin receptor and CD38 gene knockout mice

    Journal of Neuroendocrinology 22:373–379.

    https://doi.org/10.1111/j.1365-2826.2010.01976.x

    • PubMed
    • Google Scholar
54. 1. Hofer MA

    (1994) Hidden regulators in attachment, separation, and loss

    Emotion 59:192–207.

    https://doi.org/10.2307/1166146

    • Google Scholar
55. 1. Hurlemann R
    2. Patin A
    3. Onur OA
    4. Cohen MX
    5. Baumgartner T
    6. Metzler S
    7. Dziobek I
    8. Gallinat J
    9. Wagner M
    10. Maier W
    11. Kendrick KM

    (2010) Oxytocin enhances amygdala-dependent, socially reinforced learning and emotional empathy in humans

    Journal of Neuroscience 30:4999–5007.

    https://doi.org/10.1523/JNEUROSCI.5538-09.2010

    • PubMed
    • Google Scholar
56. 1. Insel TR
    2. Young LJ

    (2001) The neurobiology of attachment

    Nature Reviews Neuroscience 2:129–136.

    https://doi.org/10.1038/35053579

    • PubMed
    • Google Scholar
57. 1. Karnath HO
    2. Baier B

    (2010) Right insula for our sense of limb ownership and self-awareness of actions

    Brain Structure and Function 214:411–417.

    https://doi.org/10.1007/s00429-010-0250-4

    • PubMed
    • Google Scholar
58. 1. Keysers C
    2. Gazzola V
    3. Wagenmakers EJ

    (2020) Using Bayes factor hypothesis testing in neuroscience to establish evidence of absence

    Nature Neuroscience 23:788–799.

    https://doi.org/10.1038/s41593-020-0660-4

    • PubMed
    • Google Scholar
59. 1. Kim P
    2. Leckman JF
    3. Mayes LC
    4. Newman MA
    5. Feldman R
    6. Swain JE

    (2010) Perceived quality of maternal care in childhood and structure and function of mothers' brain

    Developmental Science 13:662–673.

    https://doi.org/10.1111/j.1467-7687.2009.00923.x

    • PubMed
    • Google Scholar
60. 1. Kim P
    2. Rigo P
    3. Leckman JF
    4. Mayes LC
    5. Cole PM
    6. Feldman R
    7. Swain JE

    (2015) A prospective longitudinal study of perceived infant outcomes at 18–24 Months: Neural and Psychological Correlates of Parental Thoughts and Actions Assessed during the First Month Postpartum

    Frontiers in Psychology 6:1–14.

    https://doi.org/10.3389/fpsyg.2015.01772

    • Google Scholar
61. 1. Kim P
    2. Strathearn L
    3. Swain JE

    (2016) The maternal brain and its plasticity in humans

    Hormones and Behavior 77:113–123.

    https://doi.org/10.1016/j.yhbeh.2015.08.001

    • PubMed
    • Google Scholar
62. 1. Kim P
    2. Capistrano CG
    3. Erhart A
    4. Gray-Schiff R
    5. Xu N

    (2017) Socioeconomic disadvantage, neural responses to infant emotions, and emotional availability among first-time new mothers

    Behavioural Brain Research 325:188–196.

    https://doi.org/10.1016/j.bbr.2017.02.001

    • PubMed
    • Google Scholar
63. 1. Kringelbach ML
    2. Lehtonen A
    3. Squire S
    4. Harvey AG
    5. Craske MG
    6. Holliday IE
    7. Green AL
    8. Aziz TZ
    9. Hansen PC
    10. Cornelissen PL
    11. Stein A

    (2008) A specific and rapid neural signature for parental instinct

    PLOS ONE 3:e1664.

    https://doi.org/10.1371/journal.pone.0001664

    • PubMed
    • Google Scholar
64. 1. Krol KM
    2. Moulder RG
    3. Lillard TS
    4. Grossmann T
    5. Connelly JJ

    (2019) Epigenetic dynamics in infancy and the impact of maternal engagement

    Science Advances 5:1–8.

    https://doi.org/10.1126/sciadv.aay0680

    • Google Scholar
65. 1. Kundakovic M
    2. Champagne FA

    (2015) Early-life experience, epigenetics, and the developing brain

    Neuropsychopharmacology 40:141–153.

    https://doi.org/10.1038/npp.2014.140

    • PubMed
    • Google Scholar
66. 1. Lancaster JL
    2. Woldorff MG
    3. Parsons LM
    4. Liotti M
    5. Freitas CS
    6. Rainey L
    7. Kochunov PV
    8. Nickerson D
    9. Mikiten SA
    10. Fox PT

    (2000) Automated talairach atlas labels for functional brain mapping

    Human Brain Mapping 10:120–131.

    https://doi.org/10.1002/1097-0193(200007)10:3<120::AID-HBM30>3.0.CO;2-8

    • PubMed
    • Google Scholar
67. 1. Leong V
    2. Byrne E
    3. Clackson K
    4. Georgieva S
    5. Lam S
    6. Wass S

    (2017) Speaker gaze increases information coupling between infant and adult brains

    PNAS 114:13290–13295.

    https://doi.org/10.1073/pnas.1702493114

    • PubMed
    • Google Scholar
68. Preprint

    1. Leong V
    2. Noreika V
    3. Clackson K
    4. Georgieva S
    5. Brightman L
    6. Nutbrown R
    7. Fujita S
    8. Neale D
    9. Wass S

    (2019) Mother-infant interpersonal neural connectivity predicts infants’ social learning

    PsyArXiv.

    https://doi.org/10.31234/osf.io/gueaq

    • Google Scholar
69. 1. Levy J
    2. Goldstein A
    3. Feldman R

    (2019) The neural development of empathy is sensitive to caregiving and early trauma

    Nature Communications 10:1905.

    https://doi.org/10.1038/s41467-019-09927-y

    • PubMed
    • Google Scholar
70. 1. Li T
    2. Chen X
    3. Mascaro J
    4. Haroon E
    5. Rilling JK

    (2017) Intranasal oxytocin, but not vasopressin, augments neural responses to toddlers in human fathers

    Hormones and Behavior 93:193–202.

    https://doi.org/10.1016/j.yhbeh.2017.01.006

    • PubMed
    • Google Scholar
71. 1. Lindquist KA
    2. Satpute AB
    3. Wager TD
    4. Weber J
    5. Barrett LF

    (2016) The brain basis of positive and negative affect: evidence from a Meta-Analysis of the human neuroimaging literature

    Cerebral Cortex 26:1910–1922.

    https://doi.org/10.1093/cercor/bhv001

    • PubMed
    • Google Scholar
72. 1. Lopatina O
    2. Inzhutova A
    3. Salmina AB
    4. Higashida H

    (2012) The roles of oxytocin and CD38 in social or parental behaviors

    Frontiers in Neuroscience 6:1–12.

    https://doi.org/10.3389/fnins.2012.00182

    • PubMed
    • Google Scholar
73. 1. Lucion AB
    2. Bortolini MC

    (2014) Mother–Pup Interactions: Rodents and Humans

    Frontiers in Endocrinology 5:1–5.

    https://doi.org/10.3389/fendo.2014.00017

    • Google Scholar
74. 1. Marlin BJ
    2. Mitre M
    3. D'amour JA
    4. Chao MV
    5. Froemke RC

    (2015) Oxytocin enables maternal behaviour by balancing cortical inhibition

    Nature 520:499–504.

    https://doi.org/10.1038/nature14402

    • PubMed
    • Google Scholar
75. 1. Martins DA
    2. Mazibuko N
    3. Zelaya F
    4. Vasilakopoulou S
    5. Loveridge J
    6. Oates A
    7. Maltezos S
    8. Mehta M
    9. Wastling S
    10. Howard M
    11. McAlonan G
    12. Murphy D
    13. Williams SCR
    14. Fotopoulou A
    15. Schuschnig U
    16. Paloyelis Y

    (2020) Effects of route of administration on oxytocin-induced changes in regional cerebral blood flow in humans

    Nature Communications 11:1–16.

    https://doi.org/10.1038/s41467-020-14845-5

    • Google Scholar
76. 1. Meaney MJ

    (2001) Maternal care, gene expression, and the transmission of individual differences in stress reactivity across generations

    Annual Review of Neuroscience 24:1161–1192.

    https://doi.org/10.1146/annurev.neuro.24.1.1161

    • PubMed
    • Google Scholar
77. 1. Naber F
    2. van Ijzendoorn MH
    3. Deschamps P
    4. van Engeland H
    5. Bakermans-Kranenburg MJ

    (2010) Intranasal oxytocin increases fathers' observed responsiveness during play with their children: a double-blind within-subject experiment

    Psychoneuroendocrinology 35:1583–1586.

    https://doi.org/10.1016/j.psyneuen.2010.04.007

    • PubMed
    • Google Scholar
78. 1. Neumann ID

    (2008) Brain oxytocin: a key regulator of emotional and social behaviours in both females and males

    Journal of Neuroendocrinology 20:858–865.

    https://doi.org/10.1111/j.1365-2826.2008.01726.x

    • PubMed
    • Google Scholar
79. 1. Neumann ID
    2. Slattery DA

    (2016) Oxytocin in general anxiety and social fear: a translational approach

    Biological Psychiatry 79:213–221.

    https://doi.org/10.1016/j.biopsych.2015.06.004

    • PubMed
    • Google Scholar
80. 1. Noriuchi M
    2. Kikuchi Y
    3. Senoo A

    (2008) The functional neuroanatomy of maternal love: mother's response to infant's attachment behaviors

    Biological Psychiatry 63:415–423.

    https://doi.org/10.1016/j.biopsych.2007.05.018

    • PubMed
    • Google Scholar
81. 1. Northoff G
    2. Heinzel A
    3. de Greck M
    4. Bermpohl F
    5. Dobrowolny H
    6. Panksepp J

    (2006) Self-referential processing in our brain--a meta-analysis of imaging studies on the self

    NeuroImage 31:440–457.

    https://doi.org/10.1016/j.neuroimage.2005.12.002

    • PubMed
    • Google Scholar
82. 1. Numan M
    2. Young LJ

    (2016) Neural mechanisms of mother-infant bonding and pair bonding: similarities, differences, and broader implications

    Hormones and Behavior 77:98–112.

    https://doi.org/10.1016/j.yhbeh.2015.05.015

    • PubMed
    • Google Scholar
83. 1. Oettl LL
    2. Ravi N
    3. Schneider M
    4. Scheller MF
    5. Schneider P
    6. Mitre M
    7. da Silva Gouveia M
    8. Froemke RC
    9. Chao MV
    10. Young WS
    11. Meyer-Lindenberg A
    12. Grinevich V
    13. Shusterman R
    14. Kelsch W

    (2016) Oxytocin enhances social recognition by modulating cortical control of early olfactory processing

    Neuron 90:609–621.

    https://doi.org/10.1016/j.neuron.2016.03.033

    • PubMed
    • Google Scholar
84. 1. Olazábal DE
    2. Young LJ

    (2006) Oxytocin receptors in the nucleus accumbens facilitate "spontaneous" maternal behavior in adult female prairie voles

    Neuroscience 141:559–568.

    https://doi.org/10.1016/j.neuroscience.2006.04.017

    • PubMed
    • Google Scholar
85. Software

    1. OSF

    (2020) OSF

    OSF.

    https://osf.io/myprojects/
86. 1. Pawluski JL
    2. Lonstein JS
    3. Fleming AS

    (2017) The neurobiology of postpartum anxiety and depression

    Trends in Neurosciences 40:106–120.

    https://doi.org/10.1016/j.tins.2016.11.009

    • PubMed
    • Google Scholar
87. 1. Peer M
    2. Salomon R
    3. Goldberg I
    4. Blanke O
    5. Arzy S

    (2015) Brain system for mental orientation in space, time, and person

    PNAS 112:11072–11077.

    https://doi.org/10.1073/pnas.1504242112

    • PubMed
    • Google Scholar
88. 1. Pratt M
    2. Zeev-Wolf M
    3. Goldstein A
    4. Feldman R

    (2019) Exposure to early and persistent maternal depression impairs the neural basis of attachment in preadolescence

    Progress in Neuro-Psychopharmacology and Biological Psychiatry 93:21–30.

    https://doi.org/10.1016/j.pnpbp.2019.03.005

    • Google Scholar
89. 1. Qin P
    2. Liu Y
    3. Shi J
    4. Wang Y
    5. Duncan N
    6. Gong Q
    7. Weng X
    8. Northoff G

    (2012) Dissociation between anterior and posterior cortical regions during self-specificity and familiarity: a combined fMRI-meta-analytic study

    Human Brain Mapping 33:154–164.

    https://doi.org/10.1002/hbm.21201

    • PubMed
    • Google Scholar
90. 1. Rassovsky Y
    2. Harwood A
    3. Zagoory-Sharon O
    4. Feldman R

    (2019) Martial arts increase oxytocin production

    Scientific Reports 9:12980.

    https://doi.org/10.1038/s41598-019-49620-0

    • PubMed
    • Google Scholar
91. 1. Riem MM
    2. Bakermans-Kranenburg MJ
    3. van IJzendoorn MH

    (2016) Intranasal administration of oxytocin modulates behavioral and amygdala responses to infant crying in females with insecure attachment representations

    Attachment & Human Development 18:213–234.

    https://doi.org/10.1080/14616734.2016.1149872

    • PubMed
    • Google Scholar
92. 1. Rilling JK
    2. Mascaro JS

    (2017) The neurobiology of fatherhood

    Current Opinion in Psychology 15:26–32.

    https://doi.org/10.1016/j.copsyc.2017.02.013

    • PubMed
    • Google Scholar
93. 1. Ross HE
    2. Cole CD
    3. Smith Y
    4. Neumann ID
    5. Landgraf R
    6. Murphy AZ
    7. Young LJ

    (2009) Characterization of the oxytocin system regulating affiliative behavior in female prairie voles

    Neuroscience 162:892–903.

    https://doi.org/10.1016/j.neuroscience.2009.05.055

    • PubMed
    • Google Scholar
94. 1. Ross HE
    2. Young LJ

    (2009) Oxytocin and the neural mechanisms regulating social cognition and affiliative behavior

    Frontiers in Neuroendocrinology 30:534–547.

    https://doi.org/10.1016/j.yfrne.2009.05.004

    • PubMed
    • Google Scholar
95. 1. Rothbart MK

    (1981) Measurement of temperament in infancy

    Child Development 52:569–578.

    https://doi.org/10.2307/1129176

    • Google Scholar
96. 1. Russell AF

    (2003) Breeding success in cooperative meerkats: effects of helper number and maternal state

    Behavioral Ecology 14:486–492.

    https://doi.org/10.1093/beheco/arg022

    • Google Scholar
97. 1. Saarimäki H
    2. Gotsopoulos A
    3. Jääskeläinen IP
    4. Lampinen J
    5. Vuilleumier P
    6. Hari R
    7. Sams M
    8. Nummenmaa L

    (2016) Discrete neural signatures of basic emotions

    Cerebral Cortex 26:2563–2573.

    https://doi.org/10.1093/cercor/bhv086

    • PubMed
    • Google Scholar
98. 1. Salomon R
    2. Malach R
    3. Lamy D

    (2009) Involvement of the intrinsic/Default system in Movement-Related self recognition

    PLOS ONE 4:e7527.

    https://doi.org/10.1371/journal.pone.0007527

    • Google Scholar
99. 1. Salomon R
    2. Levy DR
    3. Malach R

    (2014) Deconstructing the default: cortical subdivision of the default mode/intrinsic system during self-related processing

    Human Brain Mapping 35:1491–1502.

    https://doi.org/10.1002/hbm.22268

    • PubMed
    • Google Scholar
100. 1. Salomon R
     2. Ronchi R
     3. Dönz J
     4. Bello-Ruiz J
     5. Herbelin B
     6. Martet R
     7. Faivre N
     8. Schaller K
     9. Blanke O

     (2016) The insula mediates access to awareness of visual stimuli presented synchronously to the heartbeat

     The Journal of Neuroscience 36:5115–5127.

     https://doi.org/10.1523/JNEUROSCI.4262-15.2016

     • PubMed
     • Google Scholar
101. 1. Salomon R
     2. Ronchi R
     3. Dönz J
     4. Bello-Ruiz J
     5. Herbelin B
     6. Faivre N
     7. Schaller K
     8. Blanke O

     (2018) Insula mediates heartbeat related effects on visual consciousness

     Cortex 101:87–95.

     https://doi.org/10.1016/j.cortex.2018.01.005

     • PubMed
     • Google Scholar
102. 1. Schirmer A
     2. Meck WH
     3. Penney TB

     (2016) The Socio-Temporal brain: connecting people in time

     Trends in Cognitive Sciences 20:760–772.

     https://doi.org/10.1016/j.tics.2016.08.002

     • PubMed
     • Google Scholar
103. 1. Schulkin J
     2. Sterling P

     (2019) Allostasis: a Brain-Centered, predictive mode of physiological regulation

     Trends in Neurosciences 42:740–752.

     https://doi.org/10.1016/j.tins.2019.07.010

     • PubMed
     • Google Scholar
104. 1. Schultz W

     (2016) Reward functions of the basal ganglia

     Journal of Neural Transmission 123:679–693.

     https://doi.org/10.1007/s00702-016-1510-0

     • PubMed
     • Google Scholar
105. 1. Seth AK

     (2013) Interoceptive inference, emotion, and the embodied self

     Trends in Cognitive Sciences 17:565–573.

     https://doi.org/10.1016/j.tics.2013.09.007

     • PubMed
     • Google Scholar
106. 1. Seth S
     2. Lewis AJ
     3. Galbally M

     (2016) Perinatal maternal depression and cortisol function in pregnancy and the postpartum period: a systematic literature review

     BMC Pregnancy and Childbirth 16:124.

     https://doi.org/10.1186/s12884-016-0915-y

     • PubMed
     • Google Scholar
107. 1. Shamay-Tsoory SG
     2. Abu-Akel A

     (2016) The social salience hypothesis of oxytocin

     Biological Psychiatry 79:194–202.

     https://doi.org/10.1016/j.biopsych.2015.07.020

     • PubMed
     • Google Scholar
108. 1. Spreng RN
     2. Mar RA
     3. Kim AS

     (2009) The common neural basis of autobiographical memory, prospection, navigation, theory of mind, and the default mode: a quantitative meta-analysis

     Journal of Cognitive Neuroscience 21:489–510.

     https://doi.org/10.1162/jocn.2008.21029

     • PubMed
     • Google Scholar
109. 1. Sroufe LA

     (1985) Attachment classification from the perspective of infant-caregiver relationships and infant temperament

     Child Development 56:1–14.

     https://doi.org/10.2307/1130168

     • PubMed
     • Google Scholar
110. 1. Swain JE
     2. Kim P
     3. Spicer J
     4. Ho SS
     5. Dayton CJ
     6. Elmadih A
     7. Abel KM

     (2014) Approaching the biology of human parental attachment: brain imaging, oxytocin and coordinated assessments of mothers and fathers

     Brain Research 1580:78–101.

     https://doi.org/10.1016/j.brainres.2014.03.007

     • PubMed
     • Google Scholar
111. 1. Tronick EZ

     (1989) Emotions and emotional communication in infants

     American Psychologist 44:112–119.

     https://doi.org/10.1037/0003-066X.44.2.112

     • Google Scholar
112. Preprint

     1. Ulmer Yaniv A
     2. Salomon R
     3. Waidergoren S
     4. Shimon-Raz O
     5. Djalovski A
     6. Feldman R

     (2020) Synchronous caregiving from birth to adulthood tunes humans’ Social Brain

     bioRxiv.

     https://doi.org/10.1101/2020.03.09.974659

     • Google Scholar
113. 1. Valtcheva S
     2. Froemke RC

     (2019) Neuromodulation of maternal circuits by oxytocin

     Cell and Tissue Research 375:57–68.

     https://doi.org/10.1007/s00441-018-2883-1

     • Google Scholar
114. 1. Wang D
     2. Yan X
     3. Li M
     4. Ma Y

     (2017) Neural substrates underlying the effects of oxytocin: a quantitative meta-analysis of pharmaco-imaging studies

     Social Cognitive and Affective Neuroscience 12:1565–1573.

     https://doi.org/10.1093/scan/nsx085

     • Google Scholar
115. 1. Weisman O
     2. Zagoory-Sharon O
     3. Feldman R

     (2012a) Oxytocin administration to parent enhances infant physiological and behavioral readiness for social engagement

     Biological Psychiatry 72:982–989.

     https://doi.org/10.1016/j.biopsych.2012.06.011

     • PubMed
     • Google Scholar
116. 1. Weisman O
     2. Zagoory-Sharon O
     3. Feldman R

     (2012b) Intranasal oxytocin administration is reflected in human saliva

     Psychoneuroendocrinology 37:1582–1586.

     https://doi.org/10.1016/j.psyneuen.2012.02.014

     • PubMed
     • Google Scholar
117. 1. Wigton R
     2. Radua J
     3. Allen P
     4. Averbeck B
     5. Meyer-Lindenberg A
     6. McGuire P
     7. Shergill SS
     8. Fusar-Poli P

     (2015) Neurophysiological effects of acute oxytocin administration: systematic review and meta-analysis of placebo-controlled imaging studies

     Journal of Psychiatry & Neuroscience 40:E1–E22.

     https://doi.org/10.1503/jpn.130289

     • PubMed
     • Google Scholar
118. 1. Wittfoth-Schardt D
     2. Gründing J
     3. Wittfoth M
     4. Lanfermann H
     5. Heinrichs M
     6. Domes G
     7. Buchheim A
     8. Gündel H
     9. Waller C

     (2012) Oxytocin modulates neural reactivity to children's faces as a function of social salience

     Neuropsychopharmacology 37:1799–1807.

     https://doi.org/10.1038/npp.2012.47

     • PubMed
     • Google Scholar
119. 1. Yarkoni T
     2. Poldrack RA
     3. Nichols TE
     4. Van Essen DC
     5. Wager TD

     (2011) Large-scale automated synthesis of human functional neuroimaging data

     Nature Methods 8:665–670.

     https://doi.org/10.1038/nmeth.1635

     • Google Scholar
120. 1. Zink CF
     2. Meyer-Lindenberg A

     (2012) Human neuroimaging of oxytocin and vasopressin in social cognition

     Hormones and Behavior 61:400–409.

     https://doi.org/10.1016/j.yhbeh.2012.01.016

     • Google Scholar

Author details

1. Ortal Shimon-Raz

   1. IDC Herzliya, Bar Ilan University, Ramat Gan, Israel
   2. Department of Psychology, Bar Ilan University, Ramat Gan, Israel

   Contribution

   Conceptualization, Resources, Data curation, Formal analysis, Supervision, Funding acquisition, Visualization, Methodology, Writing - original draft, Project administration, Writing - review and editing

   Contributed equally with

   Roy Salomon

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-6464-8370

2. Roy Salomon

   Gonda Brain Research Center, Bar Ilan University, Ramat Gan, Israel

   Contribution

   Conceptualization, Data curation, Software, Formal analysis, Supervision, Visualization, Methodology, Writing - original draft, Project administration, Writing - review and editing

   Contributed equally with

   Ortal Shimon-Raz

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-6688-617X

3. Miki Bloch

   1. Department of Psychiatry, Tel Aviv Sourasky Medical Center, Tel Aviv, Israel
   2. Sackler Faculty of Medicine, Tel Aviv University, Tel Aviv, Israel

   Contribution

   Conceptualization, Resources, Software, Supervision, Methodology, Writing - original draft, Writing - review and editing, psychiatric evaluation of subjects

   Competing interests

   No competing interests declared

4. Gabi Aisenberg Romano

   1. Department of Psychiatry, Tel Aviv Sourasky Medical Center, Tel Aviv, Israel
   2. Sackler Faculty of Medicine, Tel Aviv University, Tel Aviv, Israel

   Contribution

   Conceptualization, Resources, Software, Visualization, Methodology, Writing - review and editing, psychiatric evaluation of subjects

   Competing interests

   No competing interests declared

5. Yaara Yeshurun

   School of Psychological Sciences, Tel Aviv University, Tel Aviv, Israel

   Contribution

   Software, Visualization, Methodology, Writing - review and editing

   Competing interests

   No competing interests declared

6. Adi Ulmer Yaniv

   1. IDC Herzliya, Bar Ilan University, Ramat Gan, Israel
   2. Gonda Brain Research Center, Bar Ilan University, Ramat Gan, Israel

   Contribution

   Software, Formal analysis, Visualization, Methodology, Writing - review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-5865-8305

7. Orna Zagoory-Sharon

   IDC Herzliya, Bar Ilan University, Ramat Gan, Israel

   Contribution

   Software, Formal analysis, Visualization, Methodology, Writing - review and editing

   Competing interests

   No competing interests declared

8. Ruth Feldman

   IDC Herzliya, Bar Ilan University, Ramat Gan, Israel

   Contribution

   Conceptualization, Resources, Software, Formal analysis, Supervision, Funding acquisition, Visualization, Methodology, Writing - original draft, Writing - review and editing

   For correspondence

   feldman.ruth@gmail.com

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0001-5048-1381

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