Neural circuits for mammalian parental behaviour

Bradley B Jamieson; Johannes Kohl

doi:10.1038/s41583-026-01073-x

. Author manuscript; available in PMC: 2026 Sep 29.

Published in final edited form as: Nat Rev Neurosci. 2026 Aug 7;27(10):687–704. doi: 10.1038/s41583-026-01073-x

Neural circuits for mammalian parental behaviour

Bradley B Jamieson ¹, Johannes Kohl ^1,^†

PMCID: PMC7619500 EMSID: EMS218257 PMID: 42567943

Abstract

Most mammalian offspring are born helpless and depend on sustained parental care for survival, making caregiving one of the most conserved and essential behavioural repertoires in mammals. Yet infant-directed behaviours can be remarkably variable, ranging from care to neglect and aggression, even within the same individual. How neural circuits support both the robustness and flexibility of parental behaviour remains poorly understood. Here, we review recent advances in our understanding of the neural mechanisms of mammalian caregiving, placing circuit-level insights from rodent models into the broader context of parental diversity across mammals. Emerging evidence indicates that internal state and social experience dynamically reshape parental circuits across timescales from hours to weeks, and that these circuits adapt their function to generate life stage-appropriate behavioural output. Rather than acting as fixed control systems for instinctive actions, parental circuits are highly plastic and context dependent. Parental behaviour therefore provides a powerful model for understanding how neural circuits are reconfigured to meet changing behavioural demands.

Introduction

Parental care is a central feature of mammalian reproduction. Compared with other vertebrates, mammals typically produce fewer offspring but invest heavily in their survival, a strategy that is particularly pronounced in species with helpless (altricial) young, such as humans, where offspring depend on prolonged caregiving^1–3. Parental behaviour comprises a stereotyped repertoire of actions. In rodents, these include retrieval of pups to the nest, grooming and nursing, as well as auxiliary behaviours such as nest building. These behaviours are expressed with remarkable consistency across individuals.

A large body of work has established that parental behaviour is mediated by dedicated neural circuits whose organisation is increasingly well understood as a result of decades of work in laboratory mice (Mus musculus), rats (Rattus norvegicus), sheep (Ovis aries) and biparental voles (Microtus ochrogaster)^4–6. Convergent evidence from these models supports the idea that there is a distributed network for parenting, spanning the extended amygdala systems involved in sensory and valence processing, hypothalamic hubs that create instinctive drive states, midbrain circuits for reward and action selection and cortical regions that contribute to learning and behavioural flexibility. In parallel, work in non-mammalian model systems, such as poison frogs and cichlids, has begun to reveal conserved principles of parental control across vertebrate classes^7–10.

Despite the apparent stereotypy of parental behaviour, a central challenge remains: parental responses to offspring are highly variable, ranging from caregiving to neglect or aggression, even within the same individual^5,11–14. This variability reflects a strong dependence of these behaviours on social experience, sensory context and internal state. Hormonal and physiological changes across timescales from hours to weeks profoundly alter parental circuit function, while prior social experience can similarly induce plasticity and alter behavioural output^5,11,15–18. Recent studies have started to uncover the cellular and circuit-level mechanisms through which these factors modulate specific components of the parental network. It is also becoming clearer how parental circuits are reconfigured throughout life. For example, a recent preprint has reported that the functional circuits supporting parental behaviour emerge earlier in development than previously appreciated, via rapid, microglia-dependent neuronal remodelling¹⁹. Together, these findings point to a high degree of plasticity in circuits traditionally viewed as hardwired and restricted to adulthood.

In this Review, we synthesise recent advances in understanding the neural basis of parental behaviour, with a focus on rodents, where most mechanistic insights have been gained. Unless otherwise stated, the discussion refers primarily to uniparental rodent species, in which maternal care predominates. Where relevant, we explicitly distinguish findings from biparental species, in which paternal care has been studied extensively. We highlight how internal state, social experience and developmental stage shape parental circuits in males and females, and discuss interactions of parental behaviour with competing motivational drives. Finally, we outline key outstanding questions and future directions for the field.

Diversity of mammalian parenting

Parental care is favoured when investment in current offspring increases lifetime fitness more than alternative uses of time and energy². In mammals, gestation and lactation create a strong asymmetry in this investment between males and females, with exclusive maternal care being the most common pattern across species^2,3. In contrast, males contribute to parenting in only 3–5% of mammals^20,21. Parental care by non-parents (alloparenting) is common, occurring in 30–50% of species. However, regular or obligate alloparenting is largely confined to cooperatively breeding species, which comprise only 10–20% of mammals^22–24. In general, whether care is provided by mothers, fathers and/or helpers is determined by a species’ ecological constraints and mating system^3,25 (Box 1).

Who provides care is a useful way to organise parental diversity. In uniparental systems, caregiving is largely restricted to mothers and tightly coupled to reproductive state^2,25. By contrast, in biparental and cooperative systems infant-directed behaviours can emerge outside pregnancy and parturition — in fathers, pair-bonded partners or non-breeding helpers — through social context and experience (Box 1). Moreover, in several biparental mammals, behavioural and endocrine changes associated with fatherhood emerge during the mate’s pregnancy, indicating that the transition into paternal care can begin before offspring are born²⁶. Cooperative breeding represents a specialised form of alloparenting in which offspring are routinely cared for by both parents and additional group members (often non-breeding helpers), illustrating that parental behaviours can be expressed independently of biological parenthood^3,27,28.

This diversity of caregiving strategies places important constraints on how parental behaviour is organised in the brain. If similar behaviours can be expressed by mothers, fathers and non-parental individuals, it is unlikely that each caregiving role relies on a wholly distinct circuit architecture. Rather, the occurrence of caregiving across parents and non-parents supports the existence of a shared neural substrate for parenting. Variation in who provides care is therefore more plausibly explained by differences in how a shared parenting circuit is recruited and modulated across physiological and social contexts than by its presence or absence. We next outline the organisation of this core parental circuit.

Core parental circuitry

Rather than being organised around a single “parenting centre”, modern circuit approaches support a distributed architecture (Fig. 1). Pro-parental circuits promote approach towards young and the execution of caregiving sequences, whereas agonistic (anti-parental) circuits suppress care or drive avoidance and attack, depending on internal state and sensory context. Cell-type- and pathway-specific manipulations, together with in vivo recordings, are increasingly revealing how this network evaluates infant cues, organises caregiving actions, and suppresses mutually incompatible behaviours.

Evaluating infant cues

A prerequisite for parenting is that infant-derived sensory cues are detected and assigned behavioural meaning. These cues are multisensory and include chemosensory, auditory, tactile and visual signals (Fig. 1)⁵. Importantly, deprivation of a single sensory modality has little effect on maternal responses, whereas disruption of two or more senses produces strong deficits, suggesting that pup recognition depends on the integration of multiple, partially redundant infant cues²⁹. In rodents, however, these modalities do not contribute equally and are sequentially engaged as the adult approaches offspring. Auditory and volatile olfactory cues are important for detecting and localising pups at a distance, whereas contact chemosensory and tactile cues become relevant during close-range investigation and direct physical interaction^30–33. Because most laboratory studies of parental behaviour in rodents are performed in darkness or low light, the contribution of vision is less well understood.

Auditory cues provide an important route by which pup distress calls are detected and translated into orienting and retrieval behaviours. In mice, pup ultrasonic vocalisations (USVs) are represented in primary auditory cortex (A1), where their encoding is strongly shaped by parental state and experience^34,35. In females, only the left A1, together with the temporal association cortex (TeA), is required for the recognition of pup vocalisations and for efficient auditory-guided retrieval^34,36,37. These auditory signals affect the hypothalamic circuitry that coordinates maternal behaviour via the posterior intralaminar thalamus (PIL), which relays information about pup calls to hypothalamic oxytocin neurons³⁸. A recent preprint has reported that a parallel A1–inferior colliculus (IC) pathway also contributes to USV-driven pup retrieval in female mice³⁹.

Chemosensation provides a second major route for infant detection and evaluation. Pup odours are detected by the main olfactory epithelium (MOE) and processed through the main olfactory bulb (OB). These cues contribute to pup localisation and approach behaviour, and disruption of MOE or OB function impairs maternal behaviour, in particular retrieval^40–43. From the OB, olfactory information is relayed to cortical and amygdalar targets, including the medial amygdala (MeA) and bed nucleus of the stria terminalis (BNST), which connect chemosensory pup representations to hypothalamic parental circuits^44–48. In sheep, bonding between ewe and lamb depends on intact olfactory processing: transient lidocaine inactivation of the MeA during the first hours after birth disrupts subsequent lamb-odour recognition, demonstrating that olfactory input to the MeA contributes to offspring-specific memory formation^49,50. In female mice, olfactory pup cues can also shape auditory pup representations in A1 through experience-dependent recruitment of basolateral amygdala (BLA) inputs carrying olfactory information from upstream olfactory cortical and amygdalar regions⁵¹. These findings suggest that olfactory and auditory pup cues are integrated in auditory cortex to support localisation of pups before direct contact.

At closer range, non-volatile contact chemosensory cues detected by the vomeronasal organ (VNO) become particularly relevant (Fig. 1)^30–33. In sexually naïve male mice, surgical or genetic ablation of the VNO abolishes pup-directed aggression and unmasks caregiving^52,53, showing that VNO input normally promotes infant-directed aggression. These cues — which include pup salivary chemosignals³² — are relayed via the accessory olfactory bulb (AOB) to the MeA, where information from the main olfactory and vomeronasal pathways can thus be integrated^54,55. In male rats, MeA lesions unmask parental care in otherwise infanticidal animals⁵⁶. It is possible that pup-directed aggression involves MeA outputs to hypothalamic aggression centres, including the anterior hypothalamic area (AHA) and ventromedial hypothalamus (VMH) (areas known to be involved in aggressive behaviours^57,58); however, direct evidence implicating these pathways specifically in pup-directed aggression is currently lacking. Nevertheless, the MeA is not simply an ‘anti-parental’ node: in males, it contains molecularly distinct populations that are associated with either pup care or male-male aggression⁵⁹. Optogenetic activation of posterodorsal MeA GABAergic neurons promotes pup grooming in female mice, whereas in males, low-intensity stimulation of these neurons similarly promotes pup grooming but stronger activation can trigger infanticide⁴⁶. Thus, chemosensory cues are likely to bias behaviour in opposite directions depending on both the identity of the MeA neuronal populations recruited and the level of their activation.

Finally, tactile cues become relevant once the adult has made direct physical contact with the infant³². Compared with auditory and chemosensory pathways, much less is known about which tactile characteristics promote caregiving (Fig. 1). In female rats, perioral anaesthesia impairs pup retrieval, and it is likely that the tactile information driving this behaviour is mediated via the infraorbital nerve^60,61. In male mice, tactile detection of physical pup features (such as limbs) contributes to pup-directed aggression, together with contact chemosensory cues³². It has recently been shown that shown that mechanosensory inputs are integrated with hypothalamic state signals in downstream areas to gate the execution of mating behaviours ⁶²; thus it is possible that a similar integration may drive parental motor actions. However, at present, the best-resolved tactile circuits are those involved in suckling and nursing. In rats, calbindin- and TIP39-expressing neurons in the PIL are activated by suckling and contribute to oxytocin and prolactin release, likely through projections to the paraventricular nucleus of the hypothalamus (PVN) and arcuate nucleus (ARC)^63–65. In addition, tyrosine hydroxylase-expressing neurons in the anteroventral periventricular nucleus (AVPV) project directly to oxytocin neurons in the PVN and are recruited during maternal interactions, providing a complementary hypothalamic pathway through which infant stimuli can promote oxytocin release and caregiving⁶⁶. PIL TIP39-expressing neurons also innervate neurons in the medial preoptic area (MPOA) of the hypothalamus that express galanin (Gal) (MPOA^Gal neurons) of the hypothalamus — a key hub within parental circuitry^47,53 — providing a pathway through which pup contact can influence central parenting circuits (Fig. 1)^64,67. At the cortical level, prolonged nursing induces plasticity in primary somatosensory cortex (S1)⁶⁸, consistent with the idea that repeated tactile interactions refine pup representation and stabilise caregiving responses over time.

How pup-derived sensory signals are represented once they reach the central parenting circuits described in the next section is poorly understood. In mother mice, concurrent pup odour and USVs, which synergistically promote approach, induce greater Fos activation in both the BNST and MPOA than either cue alone⁶⁹. More recent projection-specific studies reveal partially dissociable responses to different sensory cues in female mice: pup distress vocalisations activate lateral habenula (LHb) neurons receiving BNST input, and these neurons encode associated negative affect and are activated during pup retrieval⁷⁰, whereas projections from MPOA^Gal neurons to the hypothalamic perifornical area (PeFA) respond to live pups and pup odour, but not to pup-call playback or pup-like objects⁷¹. Thus, distinct circuit elements within the BNST and MPOA may encode different infant features.

Pro- and anti-parental central systems

Once infants have been detected, the brain must assign them behavioural meaning: should caregiving be promoted, or should it be suppressed and agonistic behaviour executed instead? A first stage of this evaluation process appears to occur within (extended) amygdala circuits. The BNST — which is strongly interconnected with the MeA, hypothalamus and olfactory regions — seems to be particularly important in converting pup-related sensory signals into motivational states. In postpartum female rats, lesions or pharmacological inhibition of the BNST disrupt pup retrieval and pup-directed care^72,73, while reversible BNST inactivation impairs both the onset and maintenance of maternal behaviour in sheep⁷⁴. Further downstream, the LHb has emerged as an important node in this circuit: pup distress vocalisations recruit BNST–LHb projections⁷⁰, and lesioning or chronically silencing the LHb in primiparous dams disrupts pup retrieval and nest building⁷⁵.

At the centre of the pro-parental network lies the MPOA, which acts as a hub for the initiation and organisation of caregiving (Fig. 1). Classical lesion studies showed that damage to the MPOA abolishes retrieval, grooming and nest building in female rats without producing gross locomotor or perceptual deficits^76,77 — findings that have since been replicated in mice^78,79 marmosets⁸⁰ and sheep⁷⁴. Conversely, electrical activation of the MPOA or local delivery of hormones such as oestradiol and prolactin can elicit maternal-like responses in virgin females and males^4,81. More recent work has refined this classical view by revealing molecularly and projection-defined MPOA subpopulations that control discrete aspects of caregiving^{14,17,18,46,47,53,70,79,82,83}.

Among these, MPOA^GAL neurons are strongly engaged during parenting; selective ablation of these neurons reduces pup-directed care, whereas activation suppresses aggression in male mice and promotes pup grooming in both sexes^47,53. Oestrogen receptor-α (Esr1)-expressing MPOA neurons (MPOA^Esr1 neurons), which make up approximately 80% of MPOA^Gal neurons⁸⁴, are particularly implicated in pup approach and retrieval, and their activity rises before action initiation^82,85. However, this population contributes to several behaviours and physiological functions, including parenting, mating and torpor^82,85,86. Another population, calcitonin receptor-expressing MPOA neurons (MPOA^Calcr neurons), are recruited during both maternal and alloparental interactions⁸⁷. In mothers, 70–85% of MPOA^Calcr neurons are Esr1-positive whereas a smaller proportion (~20%) are Gal-positive⁸⁷. Chemogenetic activation of MPOA^Calcr neurons reduces infanticide and promotes pup-directed care, whereas silencing disrupts maternal behaviour⁸⁷. The MPOA thus comprises multiple partially overlapping, molecularly defined neuronal populations that promote caregiving. The primary function of MPOA^Gal neurons — like that of other hypothalamic populations such as Esr1-expressing neurons in the ventrolateral part of the VMH (VMHvl^Esr1 neurons) — may be less to encode specific actions than to represent an affective instinctive state that biases behavioural selection toward context-appropriate actions⁸⁸.

The pro-parental neuronal populations are opposed by dedicated systems that suppress care and instead promote avoidance or pup-directed aggression. Urocortin-3-expressing neurons in the PeFA (PeFA^Ucn3 neurons) promote infant neglect and mild aggression in males and females, and receive monosynaptic inputs from stress- and parenting-associated populations — including corticotropin-releasing hormone (Crh)-expressing neurons in the PVN (PVN^Crh neurons) and MPOA^Gal neurons⁸⁹. Different PeFA^Ucn3 neuron projections seem to control distinct aspects of agonistic behaviour: whereas PeFA–VMH projections suppress pup investigation, PeFA–lateral septum (LS) stimulation promotes repulsion, and PeFA–amygdalohippocampal area (AHi) stimulation promotes pup-directed aggression⁸⁹. Consistent with this, excitatory AHi–MPOA projections drive infant-directed aggression and suppress parenting in male mice⁹⁰. Similarly, neurons in the rhomboid nucleus of the BNST (BNSTrh) have been found to be recruited during infanticide, and lesions of this structure reduce infanticidal behaviour in virgin males⁷⁹. More recent work identified a reciprocal BNST–MPOA circuit that acts as a switch between caregiving and aggression. In both male and female mice, Esr1-expressing neurons in the principal BNST (BNSTpr^Esr1 neurons) promote infant-directed aggression, whereas MPOA^Esr1 neurons promote caregiving (Fig. 1)^14,91. These two populations are coupled by mutual inhibition such that activation of one suppresses the other. Reproductive state biases behavioural output — caregiving versus aggression — by shifting the balance of activity within this antagonistic circuit (Box 2)^14,91. Where and how pro-parental and agonistic circuits communicate elsewhere in the brain remains to be shown.

Reinforcement and action

Caregiving often entails substantial energetic and opportunity costs, and competes with immediate needs such as feeding, rest and self-preservation, as discussed below. For parental behaviour to dominate action selection, infants must become strongly rewarding and infant-directed interactions must acquire strong reinforcing value. Classical operant studies in female rats showed that animals will lever-press to gain access to pups, indicating that infants themselves are powerful reinforcers⁹². Remarkably, the reinforcing value of pups, measured by lever pressing frequency, can exceed that observed for cocaine in the same paradigm, underscoring the exceptional hedonic valence of offspring cues⁹³. However, not all infant cues are uniformly attractive — pup distress vocalisations can carry aversive valence in virgin females for instance⁷⁰. How infant cues acquire positive motivational value remains an important open question. A longstanding model proposes that the MPOA acts as a parental motivational hub that enables pup stimuli to engage the mesolimbic dopamine (DA) system⁵. In this model, MPOA activity recruits ventral tegmental area (VTA) projections to the nucleus accumbens (NAc), thereby endowing infant stimuli with incentive salience and promoting pup-directed approach and caregiving.

Consistent with this framework, dopaminergic pathways centred on the VTA and NAc are central to reinforcement and performance during parenting (Fig. 1)^94,95. In postpartum rats, DA depletion reduces the responsiveness of female rats to pups and disrupts goal-directed maternal behaviours such as retrieval^96,97. Similar deficits occur following transient VTA inactivation⁹⁸. More recent work in pup-exposed virgin female mice suggests that DA signals, as in many other contexts, have reinforcement-like properties during pup interactions: the activity of VTA dopaminergic neurons (VTA^DA neurons) peaks during pup contact and then decreases with experience as retrieval becomes more efficient, and temporally precise VTA^DA neuron inhibition during pup contact decreases future retrieval performance⁸³.

Further downstream, the NAc shell appears particularly important for maternal performance. Lesions of this structure or pharmacological blockade of DA receptors therein impair pup retrieval and grooming in postpartum female rats^99,100. Moreover, individual differences in maternal behaviour correlate with stronger DA signals during mother-pup interactions¹⁰¹, consistent with a role for accumbal DA tone in sustaining maternal engagement¹⁰². Human imaging studies similarly show that infant cues recruit ventral striatal and VTA circuitry in mothers, often more strongly in response to one’s own infant than to unfamiliar infants, and that these responses correlate with measures of parent-infant attachment^103,104. Recent work in mice further indicates that cortical inputs, including orbitofrontal projections, shape VTA^DA neuron dynamics during caregiving¹⁰⁵. Together, these findings suggest that mesolimbic circuits reinforce successful infant-directed interactions, whereas hypothalamic populations such as MPOA^Gal neurons may encode the underlying parental affective or motivational state.

This parental state must then be translated into specific motor sequences. The periaqueductal gray (PAG) is a major downstream midbrain structure that contributes to the motor and autonomic implementation of caregiving as well as other instinctive responses^47,106. Lesion studies in lactating rats support a modular organisation: caudolateral and ventrolateral PAG lesions reduce suckling-induced kyphosis (a characteristic, arched-back nursing posture) and alter maternal defence (aggression towards intruders threatening the litter), whereas more rostral lesions disrupt pup retrieval^107–110. At the circuit level, projections from MPOA^Gal neurons and vesicular GABA transporter (Vgat)-expressing MPOA neurons (MPOA^Vgat neurons) to the PAG drive pup grooming and retrieval, respectively, in both sexes^47,111. In contrast, a more posterior substantia innominata (SI)–PAG pathway promotes pup-directed aggression in males¹¹². MPOA outputs may also recruit brainstem motor nuclei in the reticular formation, potentially via the retrorubral field¹¹³, but these pathways remain less well characterised.

Cortical contributions

Cortical circuits provide contextual and experience-dependent control over the largely subcortical parental network. Sensory cortex is particularly important for the representation of infant cues: as discussed above, pup vocalisations are encoded in A1 in a strongly state- and experience-dependent manner^34,114, allowing prior parental experience and oxytocin signaling to shape auditory-guided retrieval. Frontal cortical regions contribute to the flexible organisation of parental behaviour under changing environmental demands. In rats, disruption of the medial prefrontal cortex (mPFC) impairs the sequencing of maternal actions, including retrieval and nesting behaviours^115,116. In mice, mPFC neurons that project to the MPOA are preferentially engaged during maternal defence rather than caregiving, and perturbing this pathway disrupts threat-related responses¹¹⁷. Similarly, excitatory orbitofrontal cortex (OFC) projections promote experience-dependent alloparenting in virgin females, likely through modulation of VTA^DA neurons¹⁰⁵. Human imaging studies likewise implicate medial prefrontal and cingulate networks in responses to infant cues and distress^118,119. Together, these findings suggest that cortical regions refine sensory representations of offspring and enable parental behaviour to be flexibly adjusted to experience and immediate environmental context.

Parental behaviour across the lifespan

In addition to the cortical modulation of parental behaviour, substantial behavioural flexibility is already embedded within subcortical parenting circuits. Similar caregiving behaviours can be expressed by juveniles, sexually inexperienced (virgin) adults, mothers, fathers and post-reproductive individuals, indicating that functional parenting circuits are present throughout life but recruited according to behavioural need^{5,11,15,19,120,121}. Developmental stage, endocrine state and experience all act on a shared circuit scaffold to shift the balance between pro-parental and anti-parental pathways^{11,14,19,47,91}. A lifespan perspective (Fig. 2) provides a useful organising principle for understanding this parental circuit plasticity. During juvenile development in mice, rapid circuit maturation initiates caregiving towards younger siblings¹⁹; in sexually naïve adults, repeated pup exposure and social observation can promote the function of the same circuit through experience-dependent plasticity^15,35,122. Pregnancy and parturition are accompanied by profound endocrine changes that drive large-scale brain remodelling and reorganise these circuits into a stable caregiving state^11,123–125. Finally, investigations into repeated parenthood and ageing may reveal whether parental plasticity accumulates across successive reproductive experiences, and how long-lasting these changes are.

Fig. 2 — Schematic overview of life stage–dependent routes through which parental and alloparental behaviour can emerge in mice, including juvenile caregiving, experience-dependent behaviour, and parenthood. In juveniles, caregiving emerges before sexual maturity in both sexes as a developmentally regulated form of alloparental behaviour^19,128,129. In adulthood, parental behaviour can be acquired before parenthood through experience-dependent mechanisms, including repeated pup exposure (sensitisation) and observational learning from experienced caregivers^16,122,139. Parenthood is the major route into high parental performance, with pregnancy, parturition and lactation promoting maternal care, and mating, cohabitation and/or prior reproductive experience facilitating paternal behaviour in naturally biparental species and contexts in which males contribute to care^4,17,52,81. The red and blue trajectories indicate relative parental performance — i.e., the likelihood that parental behaviour is expressed and the robustness with which it is performed — in females and males, respectively, across postnatal life. Dark blue shading indicates periods during which pup-directed aggression or infanticide can occur in adult males before the transition to paternal behaviour^240–243. Approximate timing is shown for laboratory mice; this may vary in other species and/ or conditions.

Onset of alloparental behaviour in juveniles

In cooperative breeders, including humans, some non-human primates, carnivores and rodents, juveniles commonly interact with, and help care for infants, indicating that functional parental circuits predate adulthood^126,127. Juvenile alloparenting behaviour has also been observed in rats^128,129, prairie voles^130,131 and mice^120,121. A recent preprint has shown that, in laboratory mice, alloparental behaviour emerges abruptly between postnatal day 14 and 15, independently of prior social experience or ovarian hormones¹⁹. It was reported that this transition coincides with the onset of pup-responsive activity in MPOA^Gal neurons and is driven by microglia-mediated remodelling of their synaptic inputs¹⁹, which prunes an immature network into an adult-like configuration permissive for caregiving. In parallel, the maturation of the preoptic area is shaped by vomeronasal sensory inputs (Fig. 3a)¹³². Juvenile caregiving thus abruptly emerges through experience-independent remodelling of sensory inputs into parenting-controlling hypothalamic neurons.

Fig. 3 — Caregiving can emerge outside parenthood through developmental, experience-dependent and socially transmitted mechanisms. a, In juvenile mice, alloparental behaviour appears during the first month of postnatal development. Pup-derived chemosensory cues are routed through vomeronasal pathways to the medial preoptic area (MPOA), where developmental circuit maturation enables caregiving behaviour¹³². A recent preprint suggests that this transition depends on microglia-mediated remodelling of synaptic inputs onto parental MPOA neurons¹⁹. Dashed arrows indicate incompletely resolved connections or functional relationships. b, In adult virgin mice, caregiving can be acquired through repeated pup exposure (sensitisation) or observation of experienced caregivers. Sensitisation promotes caregiving, in part, through chemosensory circuits that suppress infant-directed aggression^70,145. In addition, pup vocalisations recruit oxytocin (*Oxt*) neurons in the paraventricular nucleus (PVN), and potentially the supraoptic nucleus (SON), leading to oxytocin release that promotes plasticity in sensory and motivational circuits and supports subsequent caregiving responses^16,34,38. Similarly, observation of maternal behaviour recruits oxytocinergic pathways that facilitate subsequent pup retrieval and care¹⁴⁸. Within the MPOA, oxytocin receptors (OTR) and prolactin receptors (PRLR) may provide additional routes through which neuroendocrine signals facilitate caregiving: repeated pup exposure recruits OTR-expressing neurons, whereas pup-experienced virgin females show enhanced PRLR signalling^144,147. Molecularly defined neuronal populations with established roles in parental behaviour are shown (molecular markers shown in circles). Arrows represent experimentally supported anatomical or functional relationships inferred from tracing, activity mapping and circuit perturbation studies^{16,19,34,38,47,70,83,132,138,144–146,148}. Solid lines indicate well-supported connections, whereas dashed lines denote proposed or incompletely resolved pathways. A1, primary auditory cortex; BNST, bed nucleus of the stria terminalis; CeA, central amygdala; DA, dopamine; LC, locus coeruleus; LHb,lateral habenula; PIL, posterior intralaminar thalamus; NAc, nucleus accumbens; *Oxt*, oxytocin; SC, superior colliculus; VTA, ventral tegmental area; VNO, vomeronasal organ.

Whereas both male and female juvenile mice are alloparental, infant-directed behaviours diverge in virgin adults: females often show spontaneous parental behaviour, whereas males typically exhibit pup-directed aggression⁵. This suggests that puberty reshapes parental circuits in a sex-specific manner. Indeed, it has been shown that prepubertal gonadectomy impairs subsequent parental performance in both male and female adults¹³³, and treatment of males with oestradiol or dihydrotestosterone (DHT) promotes later pup-directed aggression and alters synaptic transmission in the BNSTrh, consistent with a potentiation of anti-parental pathways¹³⁴.

Learning to care: experience-driven parenting

Repeated pup exposure can transform an initially indifferent, avoidant or even aggressive adult into a reliable caregiver, a process called sensitisation. This has been most clearly demonstrated in virgin female rodents, in which repeated pup exposure induces retrieval, licking and nursing^{15,122,135,136}. Sensitisation also occurs in males^137,138. In addition, caregiving can be acquired through observational learning, as naïve females exposed to maternal behaviour show increased responsiveness to pup cues and parental performance^34,35. Social experience can thus recruit latent caregiving circuits (Fig. 3b).

Classical studies showed that sensitisation can induce caregiving in nulliparous, ovariectomised or hypophysectomised females, indicating that this process does not require ovarian or pituitary hormones^{122,135,136,139}. Pup odours and vocalisations seem to contribute to this process^33,140,141, and — once established — these changes can persist for weeks to months, consistent with a form of maternal memory^142,143. Sensitisation seems to engage the core parental network rather than a distinct system (Fig. 3b)¹⁵. In female mice, repeated pup exposure increases activity in the MPOA¹⁴⁴ and is accompanied by enhanced oxytocin signalling: oxytocin neuron activity and peptide levels rise in the PVN and supraoptic nucleus (SON), and sensitisation is associated with increased activation of oxytocin receptor-expressing neurons in the MPOA¹⁴⁴. In line with this, artificial activation of oxytocin expressing PVN neurons (PVN^Oxt neurons) is sufficient to promote pup retrieval in virgin females¹⁴⁴. Furthermore, a recent preprint has reported that oxytocin enhances pup-call responses in A1³⁹. In parallel, sensitisation weakens aversion-related pathways, including central amygdala (CeA)–locus coeruleus (LC) projections¹⁴⁵, and BNST–LHb projections that encode negative pup valence in naïve mice⁷⁰. It also recruits reward-related circuits, engaging reinforcement-signalling VTA^DA neurons and, in males, increasing activity in the NAc, consistent with a shift in the motivational value of pups^83,138. In females, prolactin may further enhance parental responsiveness and sensitivity to pup cues^146,147.

Observational learning appears to rely on mechanisms that partially overlap with those driving sensitisation (Fig. 3b). Co-housing virgin females with dams accelerates pup retrieval, and this learning depends in part on visual observation as well as shepherding — the active guidance of a naïve individual towards pups or the nest by the dam¹⁶. Oxytocin signalling plays a central role in this process: PVN^Oxt neurons are engaged during observational learning, and both SC–PVN and PVN–A1 projections facilitate the social acquisition of parental behaviour^16,34. Strikingly, a recent preprint has shown that even viewing videos of adult conspecifics in a non-parental context is sufficient to promote parental behaviour via mechanisms that involve SC–PVN^Oxt projections, suggesting that social arousal itself can recruit latent caregiving circuits independently of direct pup experience¹⁴⁸. Comparable forms of observational learning have not been extensively studied in male mice and other rodent species. However, given the importance of social interactions in many biparental and cooperatively breeding mammals, similar mechanisms may contribute to the acquisition of caregiving beyond laboratory mice.

Reproductive experience and parenthood

Experience alone can thus recruit high levels of parental behaviour. In motherhood, the same circuitry is engaged through large-scale endocrine remodelling, stabilising caregiving as a dominant behavioural state (Fig. 4). Most mechanistic insights into the underlying neural processes come from studies in females, whereas the neural basis of paternal behaviour and fatherhood — particularly in naturally biparental species — remain far less well understood (Box 2).

Fig. 4 — a, Schematic summary of relative blood plasma hormone levels across pregnancy, parturition, lactation and weaning. Oestradiol, progesterone, prolactin and oxytocin change dynamically across this period, with levels of ovarian steroid hormones rising during pregnancy and rapidly decreasing at parturition, followed by pulsatile, suckling-associated prolactin and oxytocin release during lactation^81,165. Approximate timing shown for laboratory mice. b, These hormones act across multiple nodes of the parenting network, including the medial preoptic area (MPOA), bed nucleus of the stria terminalis (BNST), ventral tegmental area (VTA) and nucleus accumbens (NAc), to promote maternal caregiving behaviour^{4,5,11,47,82,179,203}. During lactation prolactin drives a neurotransmitter phenotype switch in arcuate (ARC) tuberoinfundibular neurons from dopamine (DA) to met-enkephalin (*Penk*) to support sustained prolactin release²⁰⁵. Other hormone-sensitive or neuromodulatory pathways that shape specific components of maternal behaviour include midbrain pathways implicated in nesting¹⁶⁴ and nursing²⁰⁰ and hypothalamic circuits that contribute to maternal aggression and defensive behaviour^187–189. Brain regions, genetically defined neuronal populations and arrows indicate experimentally supported anatomical or functional relationships inferred from tracing, activity mapping and perturbation studies^{83,151,164,169,184,187–190,197–202,205}. Solid lines indicate experimentally supported anatomical or functional relationships; dashed lines indicate proposed or incompletely resolved connections. 5HT, serotonin; *Cart*, cocaine- and amphetamine-regulated transcript; *Dat*, dopamine transporter; ESR1, oestrogen receptor alpha; EWN, Edinger-Westphal nucleus; *Gal*, galanin; MOR, mu opioid receptor; OTR, oxytocin receptor; PA, posterior amygdala; PGR, progesterone receptor; PMv, ventral premammillary nucleus; PRLR, prolactin receptor; *Vgat*, vesicular GABA transporter; VMH, ventromedial hypothalamus; *Npy2r*, Neuropeptide Y receptor 2.

Pregnancy

The transition into parenthood begins at conception. Across mammals, hormonal fluctuations ― particularly of oestradiol, progesterone, prolactin, and oxytocin ― shape physiology and behaviour during pregnancy and the postpartum period (Fig. 4). The receptors for these hormones are enriched in key nodes of parenting circuits¹²⁴. During motherhood, the brain also undergoes widespread grey matter volume changes which track hormone levels and correlate with subsequent responsiveness to infant cues and quality of mother–infant attachment (reviewed in ^123,124).

In mice, prolactin secretion increases during early pregnancy before placental lactogens become the predominant ligands for the prolactin receptor (PRLR)^149,150. Activation of hypothalamic PRLR signalling promotes infant approach and caregiving during pregnancy and has also been implicated in caregiving in males^151–153. Together, these findings indicate that lactogenic hormones act as early organisers of the parental state.

As pregnancy progresses, progesterone and oestradiol levels rise sharply (Fig. 4a), and robust parental behaviour emerges in female mice, reaching levels by late gestation that are indistinguishable from those of mothers¹¹. In rats, this transition is accompanied by increased progesterone receptor (PGR) expression in the MPOA¹⁵⁴. Classical endocrine manipulations demonstrate that these hormonal dynamics are causally linked to the onset of maternal behaviour. In ovariectomised rats, oestradiol (but not progesterone) treatment shortens the latency to pup retrieval and care, whereas prior exposure to progesterone for 4–10 days sensitises females to lower doses of oestradiol, resulting in a more robust behavioural response¹⁵⁵. Similarly, in mice, maternal behaviour is rapidly induced by oestradiol within hours of progesterone withdrawal, whereas prolonged oestradiol exposure following progesterone withdrawal impairs pup retrieval and nest building¹⁵⁶. Together, these findings indicate that the transition to maternal behaviour depends not simply on the absolute concentrations of progesterone and oestradiol, but on their temporal sequence and relative dynamics. One proposed mechanism is that sustained progesterone signalling primes MPOA neurons through PGR-dependent transcriptional and chromatin remodelling, thereby enhancing subsequent oestradiol-responsive gene expression¹⁵⁷, although the precise molecular mechanisms remain incompletely understood. Consistent with this model, coordinated late-gestation trajectories of progesterone and oestradiol predict subsequent maternal sensitivity in humans¹⁵⁸.

Recent circuit-level work has begun to explore how these hormones act on parental circuitry. Disrupting oestradiol signalling in mice impairs maternal caregiving: global or MPOA-specific ablation of Esr1 produces marked deficits in pup retrieval, nursing and grooming and can increase infanticide^82,159,160, while natural variation in MPOA Esr1 levels correlates with subsequent maternal performance^161–163. Consistent with direct hormonal action on parenting-relevant neurons, MPOA^Gal neuron-specific deletion of either Esr1 or Pgr abolishes the pregnancy-associated increase in parental behaviour and leads to persistent caregiving deficits¹¹. Oestradiol and progesterone induce complementary forms of plasticity in MPOA^Gal neurons: oestradiol reduces baseline firing while increasing neuronal excitability, whereas progesterone promotes dendritic spine formation and recruits excitatory synaptic inputs¹¹. This results in sparser MPOA^Gal neuron population activity in vivo, together with stronger, more selective responses to pup stimuli. Ovarian hormones thus remodel a central node in parental circuits during pregnancy, thereby preparing the system for caregiving. Consistent with this anticipatory role, progesterone also promotes nest building through cocaine and amphetamine regulated transcript (Cart)-expressing neurons in the Edinger–Westphal nucleus in the midbrain (Fig. 4b)¹⁶⁴.

Parturition and parenthood

Parturition marks a rapid endocrine transition that consolidates the maternal state established during pregnancy. Placental expulsion causes a sharp decline in progesterone and oestradiol levels, whereas ongoing lactogenic signalling together with oxytocin helps stabilise caregiving after parturition^66,151,165. Oxytocin release increases both peripherally, where it supports uterine contractions, and throughout the brain^166–168. In parallel, oxytocin receptor (OTR) expression is transiently upregulated in regions such as the MPOA and BNST¹⁶⁹, thereby increasing the sensitivity of parenting circuits to oxytocin. Pharmacological studies in rats and sheep show that central oxytocin administration facilitates maternal behaviour, whereas oxytocin blockade delays its onset^170–174. Conversely, genetic loss of oxytocin or OTR in mice increases the latency to initiate nurturing, although once caregiving is established these deficits are surprisingly subtle^175–179. Together, these findings indicate that oxytocin is particularly important for initiating maternal behaviour, whereas its role becomes less important once caregiving is established.

Classical studies in rats reported substantial maternal behaviour even after Caesarean delivery^180,181, indicating that pregnancy itself preconfigures much of the maternal state. However, more recent work shows that Caesarean delivery delays pup retrieval and reduces pup grooming, effects that are partially rescued by exogenous oxytocin¹⁸², suggesting that parturition-associated signals further refine parental circuit function. This transition likely depends on peripartum astrocytic remodelling, as a recent preprint reported that retraction of astrocytic processes surrounding oxytocin neurons permits greater oxytocin release in response to offspring¹⁸³.

Following birth, prolactin remains elevated to support milk production, while oxytocin is released in suckling-linked bursts to drive milk ejection¹⁸⁴. Beyond these peripheral roles, prolactin directly regulates parental circuitry: global or MPOA-specific deletion of Prlr in mice abolishes maternal care while sparing lactation^151,185.

Lactation is also accompanied by the emergence of maternal aggression, a protective component of the maternal behavioural repertoire in which mothers attack unfamiliar adult conspecifics to protect their offspring¹⁸⁶. Maternal aggression is mediated by a distributed network that includes the posterior amygdala (PA), VMHvl and ventral premammillary nucleus (PMv)^187,188. Recent work identified an excitatory projection from Esr1-expressing neurons in PA (PA^Esr1) neurons to neuropeptide Y receptor 2-expressing neurons in VMHvl (VMHvl^Npy2r) neurons as a key component of maternal aggression circuitry in mice¹⁸⁸. During lactation, both PA–VMHvl synaptic transmission and VMHvl^Npy2r neuron excitability are enhanced, while oxytocin signalling increases the responsiveness of PA^Esr1 neurons, collectively promoting robust maternal defence¹⁸⁸. Prolactin exerts multiple, region-specific effects within these circuits. In the VMH, prolactin signalling restrains maternal aggression through Prlr-expressing neurons¹⁸⁹. In parallel, prolactin and oxytocin transiently recruit a previously quiescent population of dopamine transporter-expressing PMv (PMv^Dat) neurons, increasing their excitability and enabling the expression of maternal aggression at the expense of ongoing pup retrieval or nursing (Fig. 4b)¹⁸⁷.

Oxytocin and vasopressin (AVP) provide additional neuromodulatory control across distributed sensory, hypothalamic, limbic and reward circuits to support maternal performance after birth. In mice, oxytocin signalling in the VTA enhances DA release in the NAc and correlates with individual differences in maternal grooming¹⁹⁰. AVP exerts a parallel modulatory influence on parental behaviour: in rats, central AVP administration enhances caregiving, whereas blockade of its receptors — particularly V1a (AVPR1A) and V1b (AVPR1B) — reduces both maternal care and maternal aggression^191–195. Consistent with this, genetic deletion of Avpr1a impairs maternal behaviour in Syrian hamsters (Mesocricetus auratus)¹⁹⁶.

Monoamines and endogenous opioids provide further state-dependent modulation of parental circuits. In rats, DA release in the NAc correlates with maternal licking and pup grooming¹⁰¹, and blocking accumbal DA receptors disrupts caregiving⁹⁹, in part via oxytocin-sensitive VTA–NAc projections¹⁹⁰. Noradrenergic LC–anterior cingulate cortex (ACC) signalling supports maternal behaviour in mice^197,198, while serotonin (5-HT) released from the rat dorsal raphe (DR) is required for nursing through actions on 5HT2A receptors in the mPFC rather than directly within the MPOA^199–201. By contrast, endogenous opioid signalling appears to constrain caregiving: in rats, hypothalamic μ-opioid receptor expression decreases after parturition²⁰², morphine suppresses maternal behaviour²⁰³, and opioid receptor antagonism enhances nursing²⁰⁴. Notably, tuberoinfundibular DA neurons in the ARC may switch to enkephalin release under high-prolactin conditions in mice, providing a potential mechanism by which endocrine state engages opioid modulation of parental circuits²⁰⁵. Several layers of hormonal and neuromodulatory control thus act together to promote the onset and stabilisation of robust caregiving behaviour during parenthood.

Long-term plasticity and parental experience

Reproductive experience can have enduring effects on parental behaviour and its underlying neural substrates. Prior pregnancy and caregiving accelerate the onset of maternal responses in subsequent reproductive cycles and can produce lasting changes in defensive components of care^206,207. More broadly, pregnancy and motherhood induce persistent physiological and neural adaptations, although it remains unclear to what extent later maternal behaviour depends on retained circuit modifications versus endocrine changes occurring during each new pregnancy and parturition^123,208,209.

Some of these adaptations persist well beyond the reproductive period itself. In rodents, parity is associated with lasting cellular and molecular brain changes²¹⁰. In humans, childbirth is linked to persistent structural alterations, including a younger-looking brain-age profile in midlife²¹¹, and longitudinal imaging studies show sustained changes in grey matter volume, cortical thickness and white matter microstructure following pregnancy²¹². Together, these findings suggest that parenthood can leave lasting imprints on the brain.

In some species, these persistent changes may support continued caregiving roles beyond reproduction, although this has not been experimentally tested. In killer whales (Orcinus orca), post-reproductive females enhance grandoffspring survival and assume leadership roles during periods of resource scarcity^213,214. Similarly, in Asian elephants (Elephas maximus), the presence of a grandmother improves calf survival and maternal reproductive success in daughters²¹⁵. In humans, too, grandmaternal investment is associated with beneficial effects on descendant outcomes²¹⁶, although these effects likely reflect a combination of neural, social and ecological mechanisms.

Balancing care with competing drives

Parenting is costly, can increase exposure to danger, and — in some contexts — must be balanced with sexual behaviour to maximise lifetime reproductive success. Even in experienced parents, infant-directed behaviour is therefore continuously balanced against other urgent behavioural demands, including feeding, threat and stress avoidance, mating and rest. A central challenge is thus to understand how parental circuits communicate with other neural systems to organise this behavioural competition.

One of the clearest examples is feeding^12,217,218. In mice, the presence of pups can suppress hunger-driven food intake²¹⁹, and activation of excitatory MPOA neurons is sufficient to inhibit feeding²¹⁹. This competition is bidirectional: food deprivation or activation of agouti-related peptide (AGRP) neurons in the ARC — which orchestrate the behavioural effects of hunger — can promote pup neglect or pup-directed or aggression in virgin female mice by inhibiting MPOA neurons through the release of GABA and neuropeptide Y (NPY)^12,217. Importantly, both hormonal state and social experience shape this interaction. Fluctuations in progesterone-to-oestradiol hormone ratio across the oestrous cycle alter neuronal excitability in the MPOA, thereby altering the probability that hunger-evoked NPY release will suppress MPOA activity and promote pup-directed aggression¹². In contrast, food deprivation does not induce pup-directed aggression in pup-sensitised virgin females^12,217, indicating that prior social experience can stabilise caregiving in the face of metabolic challenge. Beyond the hypothalamus, fasting also increases NPY signalling through Y1 receptors in the mouse dorsal raphe nucleus, which contributes to the suppression of maternal care²²⁰. Together, these findings indicate that the balance between care and feeding is dynamically shaped by energetic need, endocrine state and prior social experience.

Caregiving must also be balanced against defensive responses to threat and stress. Stress can strongly disrupt parental behaviour, reducing pup retrieval, grooming and nursing while increasing avoidance, neglect and, in extreme cases, infant-directed aggression in female rats and mice^221–223. Anti-parental neuronal populations such as PeFA^Ucn3 neurons provide a route through which stress signals can suppress caregiving: these neurons receive input from both stress- and parenting-associated neuronal populations, including Crh-expressing neurons in the PVN and MPOA^Gal neurons, and their activation promotes infant neglect and aggression⁸⁹. Direct manipulation of CRH signalling similarly suppresses parental behaviour. In ovariectomized, steroid-primed virgin female rats, intracerebroventricular CRH administration suppresses pup retrieval and grooming and, at higher doses, induces infanticide²²⁴. These effects are anatomically specific: CRH signalling in the MPOA impairs retrieval and especially nursing²²⁵, whereas in the anterior-dorsal BNST it induces maternal neglect²²⁶. Here, too, prior experience reduces both the magnitude and duration of this inhibition and prevents CRH-induced infanticide²²⁴. Motherhood provides an additional layer of resilience to stress. Parental behaviour in dams is remarkably resistant to perinatal stress and corticosterone treatment²²⁷, and defensive responses are reorganised to prioritize offspring protection: when exposed to threat, dams preferentially run toward an exposed pup and carry it to safety instead of fleeing, and this seems to be driven by mPFC projections to MPOA^Esr1 neurons¹¹⁷.

A third dimension of behavioural competition is between parental care and mating. This is particularly evident during postpartum oestrus, when females can be simultaneously maternally and sexually motivated, and the relative expression of these behaviours depends on pup age, hormonal state and environmental context^5,228. Although the VMHvl is considered the principal hypothalamic regulator of female sexual receptivity^229–232, activation of MPOA neurons suppresses female sexual motivation after mating, suggesting that the MPOA may also contribute to coordinating parental and sexual motivational states²³³. Although little is currently known about local connectivity within the MPOA, local microcircuit organisation might contribute to switching between these competing behavioural states. Recent work also suggests that, in mice, there is a partial overlap between MPOA neurons associated with parenting and prosocial behaviour (helping behaviour directed at adult conspecifics)²³⁴, thus lending support to the longstanding hypothesis that prosocial behaviours build on neural systems originally evolved for parental care²³⁵. Similarly, in marmosets, partial overlap between parenting- and mating-associated MPOA neurons suggests that this organisation may reflect a conserved principle of shared hypothalamic control over social behaviour⁸⁰.

More generally, parental circuits are shaped by numerous internal drives and motivational states, including hunger, stress, arousal, sexual motivation, sleep pressure and reproductive state. Where and how this high-dimensional state space is integrated in the brain and translated into context-appropriate social actions remains largely unresolved and represents one of the major outstanding challenges in understanding parenting and behaviour in general.

Conclusions and perspectives

Recent work has substantially advanced our understanding of the neural basis of parental behaviour. Across life stages, a common, highly distributed circuit scaffold can be recruited in juveniles, non-parental adults, mothers and fathers, and may also support caregiving by post-reproductive individuals in some species. This circuit scaffold is profoundly shaped by endocrine state, social experience, developmental stage and competing motivational demands. Parental circuits are therefore both robust and flexible: robust in that they can generate caregiving with little or no prior experience, and flexible in that their output is continuously recalibrated according to internal state and external demands.

A central conceptual question is whether parental behaviour is controlled by a dedicated and molecularly distinct circuit, or whether it instead emerges from more general-purpose social-behavioural circuits shared with mating, aggression and prosociality. Recent evidence for overlap between parenting-related and other hypothalamic populations argues against a simple one-circuit-one-behaviour view²³⁴. In this framework, behavioural specificity may not arise from fully segregated circuit modules, but from how shared circuit elements are differentially recruited by sensory context and internal state. This question extends to what should count as part of a ‘parental circuit’: should inclusion require selective activity during parenting and necessity or sufficiency for caregiving, or is a modulatory role enough? This is particularly relevant for cortical and neuromodulatory systems, which may not be parenting-specific but clearly shape parental performance under specific conditions. Resolving these questions will require longitudinal and cell-type-specific recordings across behavioural contexts, combined with improved activity-labelling approaches that can directly test whether the same neurons are reused across parenting, mating and aggression.

A second major challenge is understanding how parental states are translated into specific actions. Considerable progress has been made in identifying hypothalamic and limbic nodes that promote or suppress caregiving, but far less is known about how these states are converted into motor programmes—such as retrieval, crouching, nest building, licking, defence or attack—through downstream midbrain and brainstem circuits. As recently shown for mating, these transformations may rely on downstream gating mechanisms that integrate hypothalamic state signals with sensory inputs to permit specific motor actions⁶². Determining these output pathways will be essential for understanding how hypothalamic state signals are converted into adaptive behavioural sequences.

The neural basis of paternal behaviour remains far less well understood than that of motherhood, particularly the post-mating switch from infanticide to care and the hormonal and circuit changes associated with fatherhood. Likewise, we know surprisingly little about how parental circuitry differs across species with uniparental, biparental or cooperative breeding systems. Do these species rely on modulation of a conserved circuit architecture, or do they differ in more fundamental aspects of functional connectivity and circuit composition? Work in voles and oldfield mice suggests that species differences in parental strategy can arise through differential neuromodulation of a shared circuit architecture rather than fundamentally distinct circuitry^236,237, but whether this generalises across mammals remains unclear. Finally, understanding what fails when parental behaviour is disrupted — for example in postpartum psychiatric disorders — remains an important translational challenge. Defining how parental circuits become unstable or maladaptive may provide broader insight into how social behaviour breaks down in disease.

Addressing these questions will require both technical and conceptual advances. In established model systems, especially mice, increasingly powerful tools for longitudinal and cell-type-specific neural recording, improved genetic access to developmentally defined circuit elements, and better approaches to monitor and manipulate neural plasticity should make it possible to track how parental circuits emerge and change over time. At the same time, studying parenting in more naturalistic conditions will be crucial, as many key features of caregiving — including social structure, offspring demand, ecological constraints and competing motivational states — are difficult to capture in simplified laboratory assays. Comparative work in emerging model systems, particularly species that exhibit cooperative breeding, will be equally important for establishing conserved principles of parental circuit organisation^80,236,238.

Parenting has long served as a powerful model for studying instinctive behaviour. It is now becoming equally valuable for understanding how neural circuits balance stability with flexibility, and how they are remodelled to meet ever-changing behavioural demands across physiological states and throughout life. Understanding the parental brain will therefore likely reveal broader principles by which the nervous system generates adaptive social behaviour.

Box 1. Mammalian parenting strategies.

Sex differences in mammalian parental care emerge from asymmetries in the reproductive investment of males and females and from ecological conditions that shape the relative fitness payoffs of mating versus caregiving². Because mammalian females necessarily invest in gestation and lactation, their minimum required parental investment is typically higher than that of males, biasing the balance toward maternal care^2,3. Biparental care or cooperative breeding evolves when the involvement of additional caregivers substantially increase offspring survival and when the costs of care are reduced. In the case of alloparental care, this occurs when the marginal fitness benefits of additional care are high, such as when offspring are numerous, highly dependent or costly to provision, carry or protect, and when the costs to caregivers are sufficiently low or offset by direct or indirect fitness gains. Both biparental care and cooperative breeding are often associated with social monogamy, which tends to evolve when breeding females are spatially dispersed and intolerant of one another, limiting a male’s ability to monopolise multiple mates; under these conditions, guarding a single female becomes an efficient strategy^23,25.

Biparental or paternal care (Box 2) often follows the evolution of social monogamy, consistent with the idea that sustained male–female association creates conditions in which male care becomes advantageous²³. Prominent examples of this type of care include California mice (Peromyscus californicus), which demonstrate biparental care, and prairie voles (Microtus ochrogaster), which exhibit substantial paternal care compared with uniparental rats and mice^244,245. Recent work in monogamous oldfield mice (Peromyscus polionotus has demonstrated one mechanism involved in the evolution of socially monogamous parental behaviour. Oldfield mice have evolved a novel adrenal cell type that produces 20α-hydroxyprogesterone, a progesterone metabolite that promotes parental care after conversion to allo-diol in the brain, where it acts as a GABA_A receptor antagonist²³⁶.

Cooperative breeding — in which offspring are routinely cared for by individuals in addition to their parents — has evolved in a small number of socially monogamous species that regularly produce multiple offspring per litter, including meerkats (Suricata suricatta), marmosets (Callithrix jacchus) and tamarins (Saguinus oedipus)^27,246–248. In these species, the combination of stable pair bonds and high offspring demand likely increases the marginal benefit of additional caregivers: multiple dependent offspring substantially raise provisioning and carrying demands, while the existence of a stable breeding pair can promote delayed dispersal of older offspring, creating opportunities for helping behaviour²⁷. In such species, multiple group members contribute to protecting, warming, feeding and/or carrying infants, and these contributions are associated with enhanced offspring fitness^246,247,249.

Box 2. Paternal care.

Paternal care provides an opportunity to understand how parenting circuits can be recruited independently of pregnancy. Although rare across mammals, comparative phylogenetic analyses suggest that biparental care has evolved repeatedly from female-only care²⁵⁰. At the same time, paternal care often co-occurs with pair bonding, territorial defence and changes in social status (for example, becoming a resident, pair-bonded breeding male), complicating efforts to distinguish mechanisms specific to caregiving^20,251. Fathers in biparental species perform many of the same non-lactational pup-care behaviours as mothers^252,253, including retrieving, grooming and huddling over pups^252,253. Within Peromyscus, paternal care is observed in monogamous oldfield and California mice, but is largely absent from promiscuous eastern deer mice (Peromyscus maniculatus) and white-footed mice (Peromyscus leucopus)^254,255.

No pregnancy-like endocrine ‘on-switch’ for parental behaviour has been identified in males. Instead, studies in both wild and laboratory mice showed that ejaculation triggers a delayed post-mating switch, such that pup-directed attack is progressively suppressed and parental responses emerge around the time the male’s own offspring would be expected^240–243. However, mating alone is not always sufficient. Cohabitation with a pregnant mate can suppress infanticide even before birth, whereas males housed with non-pregnant females remain far more likely to attack pups^256,257. The post-mating switch is reversible, but experienced fathers show a more stable suppression of infant-directed aggression regardless of the female’s current reproductive state^256,257. Thus, paternal behaviour is regulated by a combination of post-mating timing, social context and prior experience. The relative contributions of copulation, cohabitation and female reproductive state differ across genetic backgrounds^52,258,259. Work in African striped mice (Rhabdomys pumilio) further shows that prior mating is not an absolute prerequisite for male care and that the MPOA expression level of the Agouti gene scales with group density and is negatively associated with parental behaviour²³⁸.

In mice, the male transition to caregiving seems to occur in two stages: from infanticide to pup tolerance, and from tolerance to active caregiving. Ejaculation appears sufficient to suppress aggression, whereas cohabitation with a pregnant female and subsequent pup exposure promote the emergence of full paternal behaviour^14,91. Distinct forms of circuit plasticity seem to support these transitions. Mating and cohabitation reduce inhibition from cocaine- and amphetamine-regulated transcript (CART)-positive medial amygdala neurons onto central medial preoptic area (MPOA) neurons, whereas parental experience strengthens inhibitory transmission onto neurons in the rhomboid nucleus of the bed nucleus of the stria terminalis (BNSTrh), which drive pup-directed aggression in both males and females^14,91. These findings support a model in which different nodes of the parenting circuit are sequentially remodelled to first suppress infanticide and then promote active care.

At the circuit level, paternal behaviour recruits much of the same architecture that supports maternal care. Pup-directed aggression strongly depends on vomeronasal and tactile pathways responsive to both pup-derived chemosignals and physical pup features^32,52. As in females, MPOA lesions switch fathers from caregiving to infanticide, whereas BNSTrh lesions reduce infanticide in virgin males⁷⁹. Similarly in naturally biparental California mice, MPOA lesions disrupt parental behaviour in both males and females^260,261. Additional shared anti-parental pathways include amygdalohippocampal area (AHi)– MPOA projections, which promote aggression over caregiving, and UCN3-expressing perifornical area (PeFA) neurons, which drive infant-directed neglect and aggression^89,90. A recent preprint has suggested that reciprocal MPOA–BNST dynamics are particularly important⁹¹. Recent work further identified a CART- and ESR1-expressing population of neurons in the principal BNST that drives infant-directed aggression: fathers show strengthened inhibitory synaptic input onto these neurons from the anterior commissure nucleus (ACN)²⁶².

Hypothalamic oxytocin and arginine vasopressin (AVP) signalling play a central role in paternal caregiving. In male mice, oxytocin release from PVN neurons is required for paternal behaviour, and optogenetic activation of these neurons is sufficient to induce caregiving even in virgin males¹⁷. During fatherhood, excitatory input from the lateral hypothalamus onto oxytocin-expressing PVN neurons (PVN^Oxt neurons) is strengthened, enhancing oxytocin release and promoting caregiving¹⁷. AVP-expressing PVN neurons also promote caregiving, acting via AVP-sensitive oxytocin receptor signalling in the MPOA and ACN²⁶³. Similar mechanisms likely exist in other species as PVN^Oxt neurons are also activated during paternal behaviour and promote caregiving in male mandarin voles (Lasiopodomys mandarinus)²⁶⁴.

Endocrine and neuromodulatory influences on paternal care are generally subtler than in females. In California mice, castration reduces paternal care and pup grooming, whereas testosterone (T) replacement restores both²⁶⁵. This effect may involve local aromatase signalling within the MPOA, and transient T rises during courtship can predict later paternal behaviour^266,267. In prairie voles, castration reduces paternal responsiveness and AVP-positive projections to the lateral septum (LS)²⁶⁸, although paternal care can remain relatively steroid-independent in other species, including Campbell’s dwarf hamsters (Phodopus campbelli)^269,270. In humans, T typically declines across the transition to fatherhood and lower T is more consistently associated with caregiving and relationship investment than with mating effort^271,272. In parallel, prolactin and oxytocin levels respond to infant cues and parent–infant contact in both humans and rodents^273–276 and basal circulating prolactin is required for paternal responses in male mice¹⁵³. Collectively, current evidence supports a model in which paternal care emerges primarily through state-dependent recruitment and sequential remodelling of a shared caregiving circuit rather than via male-specific pathways^47,85.

Acknowledgements

We thank members of the State-dependent Neural Processing Lab for discussions and comments on the manuscript.

Glossary

Agonistic behaviour: Any social behaviour related to fighting, conflict, or competition, including threats, aggression, and submission.
Alloparenting: Care of offspring provided by individuals other than biological parents, including extended family or non-kin.
Valence: The positive or negative significance assigned to a stimulus, which biases behaviour towards approach or avoidance.
Cooperative breeding: A social system in which, in addition to the breeding parents, non-breeding group members consistently help care for offspring.
Core parental circuit: The conserved set of neural populations and pathways that organises infant-directed caregiving behaviour.
Distributed architecture: A mode of neural organization in which behaviour emerges from interactions across multiple brain regions rather than from a single control centre.
Gonadectomy: Surgical removal of the gonads (ovaries or testes), used experimentally to eliminate endogenous sex hormone production. The term gonadectomised refers to individuals that have undergone this procedure.
Hypophysectomised: Refers to individuals that have undergone surgical removal of the pituitary gland, resulting in loss of pituitary hormone secretion.
Internal state: The physiological or motivational condition of an animal, including hormonal, metabolic or affective factors, that shapes how sensory cues are processed and translated into behaviour.
Nulliparous: Refers to an individual that has never given birth. Commonly used to describe females without prior reproductive experience.
Observational learning: The acquisition of behaviour through observing another individual perform that behaviour.
Parturition: The process of giving birth, marking the transition from pregnancy to the postpartum state.
Peripartum: The period immediately before, during and after parturition.
Reproductive state: The current physiological status of an individual in relation to reproduction, for example virgin, pregnant, lactating or postpartum.
Sensitisation: The gradual emergence of parental behaviour in a non-parental animal following repeated exposure to infants.
Social monogamy: A social system in which one male and one female form a stable pair association, often associated with joint territory defence or offspring care.
Vomeronasal organ (VNO): A chemosensory organ that detects non-volatile social cues and contributes importantly to close-range evaluation of conspecifics.

Footnotes

Author Contributions

B.B.J. and J.K. conceptualised the manuscript, prepared the figures, wrote the original draft of the manuscript, and reviewed and edited the manuscript.

Competing interests

The authors declare no competing interests.

References

1.Stearns SC. The Evolution of Life Histories. Oxford University Press; Oxford, New York: 1992. [Google Scholar]
2.Trivers RL. In: Foundations of Animal Behavior: Classic Papers with Commentaries. Houck LD, Drickamer LC, editors. University of Chicago Press; Chicago, IL: 1996. Parental Investment and Sexual Selection. [Google Scholar]
3.Clutton-Brock TH. The Evolution of Parental Care. Princeton University Press; Princeton, N.J: 1991. [Google Scholar]
4.Numan M, Insel T. The Neurobiology of Parental Behavior. Springer New York; New York, NY: 2003. [DOI] [Google Scholar]
5.Numan M. The Parental Brain: Mechanisms, Development, and Evolution. Oxford University Press; New York: 2020. [DOI] [Google Scholar]
6.Kaplan HS, Horvath PM, Rahman MM, Dulac C. The neurobiology of parenting and infant-evoked aggression. Physiol Rev. 2025;105:315–381. doi: 10.1152/physrev.00036.2023. [DOI] [PMC free article] [PubMed] [Google Scholar]
7.Maciejewski MF, Bell AM. Insights into Parental Care from Studies on Non-mammalian Vertebrates. Affect Sci. 2022;3:792–798. doi: 10.1007/s42761-022-00127-4. [DOI] [PMC free article] [PubMed] [Google Scholar]
8.Fischer EK, et al. The neural basis of tadpole transport in poison frogs. Proc Biol Sci. 2019;286:20191084. doi: 10.1098/rspb.2019.1084. [DOI] [PMC free article] [PubMed] [Google Scholar]
9.Roland AB, O’Connell LA. Poison frogs as a model system for studying the neurobiology of parental care. Curr Opin Behav Sci. 2015;6:76–81. [Google Scholar]
10.Cunha-Saraiva F, et al. Parental care and neuropeptide dynamics in a cichlid fish Neolamprologus caudopunctatus. Horm Behav. 2019;116:104576. doi: 10.1016/j.yhbeh.2019.104576. [DOI] [PubMed] [Google Scholar]
11.Ammari R, et al. Hormone-mediated neural remodeling orchestrates parenting onset during pregnancy. Science. 2023;382:76–81. doi: 10.1126/science.adi0576. [ This study finds that remodelling of Galanin-expressing MPOA neurons by the ovarian hormones oestradiol and progesterone is critical for the onset of parental behaviour during pregnancy in mice ] [DOI] [PMC free article] [PubMed] [Google Scholar]
12.Cao M, et al. Integration of hunger and hormonal state gates infant-directed aggression. Nature. 2025;648:138–145. doi: 10.1038/s41586-025-09651-2. [DOI] [PMC free article] [PubMed] [Google Scholar]
13.Miyamichi K. Neural basis for behavioral plasticity during the parental life-stage transition in mice. Front Neural Circuits. 2024;17:1340497. doi: 10.3389/fncir.2023.1340497. [DOI] [PMC free article] [PubMed] [Google Scholar]
14.Mei L, Yan R, Yin L, Sullivan RM, Lin D. Antagonistic circuits mediating infanticide and maternal care in female mice. Nature. 2023;618:1006–1016. doi: 10.1038/s41586-023-06147-9. [ This study establishes that reciprocal inhibition between oestrogen receptor α (ESR1)-expressing cells in the MPOA and BNST control the balance between pup-directed care vs aggression in female mice ] [DOI] [PMC free article] [PubMed] [Google Scholar]
15.Stolzenberg DS, Champagne FA. Hormonal and non-hormonal bases of maternal behavior: The role of experience and epigenetic mechanisms. Horm Behav. 2016;77:204–210. doi: 10.1016/j.yhbeh.2015.07.005. [DOI] [PubMed] [Google Scholar]
16.Carcea I, et al. Oxytocin neurons enable social transmission of maternal behaviour. Nature. 2021;596:553–557. doi: 10.1038/s41586-021-03814-7. [ This study identifies oxytocin neurons as a neural substrate for the social transmission of maternal behaviour through experience-dependent cortical plasticity ] [DOI] [PMC free article] [PubMed] [Google Scholar]
17.Inada K, et al. Plasticity of neural connections underlying oxytocin-mediated parental behaviors of male mice. Neuron. 2022;110:2009–2023.:e5. doi: 10.1016/j.neuron.2022.03.033. [ This study demonstrates that fatherhood remodels long-range synaptic inputs onto hypothalamic oxytocin neurons, enabling the transition from pup-directed aggression to paternal caregiving in male mice ] [DOI] [PubMed] [Google Scholar]
18.Stagkourakis S, et al. A Neuro-hormonal Circuit for Paternal Behavior Controlled by a Hypothalamic Network Oscillation. Cell. 2020;182:960–975.:e15. doi: 10.1016/j.cell.2020.07.007. [ This elegant study identifies distinct electrical oscillation patterns in neuroendocrine dopamine neurons as the basis of species-specific paternal strategies in male mice and rats ] [DOI] [PMC free article] [PubMed] [Google Scholar]
19.Jamieson BB, et al. Overnight circuit remodelling drives juvenile alloparental care. 2025 doi: 10.1101/2025.06.13.659524. Preprint at. [DOI] [Google Scholar]
20.Rogers FD, Bales KL. Mothers, Fathers, and Others: Neural Substrates of Parental Care. Trends Neurosci. 2019;42:552–562. doi: 10.1016/j.tins.2019.05.008. [DOI] [PMC free article] [PubMed] [Google Scholar]
21.Rilling JK, Mascaro JS. The neurobiology of fatherhood. Curr Opin Psychol. 2017;15:26–32. doi: 10.1016/j.copsyc.2017.02.013. [DOI] [PubMed] [Google Scholar]
22.Clutton-Brock T. Cooperation between non-kin in animal societies. Nature. 2009;462:51–57. doi: 10.1038/nature08366. [DOI] [PubMed] [Google Scholar]
23.Lukas D, Clutton-Brock TH. The Evolution of Social Monogamy in Mammals. Science. 2013;341:526–530. doi: 10.1126/science.1238677. [DOI] [PubMed] [Google Scholar]
24.Hrdy SB. Mothers and Others: The Evolutionary Origins of Mutual Understanding. Vol. 422 Harvard University Press; Cambridge, MA, US: 2009. [Google Scholar]
25.Kleiman DG. Monogamy in Mammals. Q Rev Biol. 1977;52:39–69. doi: 10.1086/409721. [DOI] [PubMed] [Google Scholar]
26.Saltzman W, et al. Paternal Care in Biparental Rodents: Intra- and Inter-individual Variation. Integr Comp Biol. 2017;57:589–602. doi: 10.1093/icb/icx047. [DOI] [PMC free article] [PubMed] [Google Scholar]
27.Lukas D, Clutton-Brock T. Climate and the distribution of cooperative breeding in mammals. R Soc Open Sci. 2017;4:160897. doi: 10.1098/rsos.160897. [DOI] [PMC free article] [PubMed] [Google Scholar]
28.Hrdy SB, Burkart JM. The emergence of emotionally modern humans: implications for language and learning. Philos Trans R Soc B Biol Sci. 2020;375:20190499. doi: 10.1098/rstb.2019.0499. [DOI] [PMC free article] [PubMed] [Google Scholar]
29.Beach FA, Jaynes J. Studies of maternal retrieving in rats. III. Sensory cues involved in the lactating female’s response to her young. Behaviour. 1956;10:104–125. [Google Scholar]
30.Rymer TL. The Role of Olfactory Genes in the Expression of Rodent Paternal Care Behavior. Genes. 2020;11:292. doi: 10.3390/genes11030292. [DOI] [PMC free article] [PubMed] [Google Scholar]
31.Kohl J, Autry AE, Dulac C. The neurobiology of parenting: A neural circuit perspective. BioEssays. 2017;39:1–11. doi: 10.1002/bies.201600159. [DOI] [PMC free article] [PubMed] [Google Scholar]
32.Isogai Y, et al. Multisensory Logic of Infant-Directed Aggression by Males. Cell. 2018;175:1827–1841.:e17. doi: 10.1016/j.cell.2018.11.032. [ This study finds that combined detection of chemosensory and tactile cues elicits pup-directed aggression in male mice ] [DOI] [PMC free article] [PubMed] [Google Scholar]
33.Noirot E. Selective priming of maternal responses by auditory and olfactory cues from mouse pups. Dev Psychobiol. 1969;2:273–276. doi: 10.1002/dev.420020413. [DOI] [PubMed] [Google Scholar]
34.Marlin BJ, Mitre M, D’amour JA, Chao MV, Froemke RC. Oxytocin enables maternal behaviour by balancing cortical inhibition. Nature. 2015;520:499–504. doi: 10.1038/nature14402. [DOI] [PMC free article] [PubMed] [Google Scholar]
35.Schiavo JK, et al. Innate and plastic mechanisms for maternal behaviour in auditory cortex. Nature. 2020;587:426–431. doi: 10.1038/s41586-020-2807-6. [ This study finds that experience-dependent maternal behaviour in mice results from interactions between intrinsic mechanisms and experience-dependent plasticity in the auditory cortex ] [DOI] [PMC free article] [PubMed] [Google Scholar]
36.Tasaka G, et al. Genetic tagging of active neurons in auditory cortex reveals maternal plasticity of coding ultrasonic vocalizations. Nat Commun. 2018;9:871. doi: 10.1038/s41467-018-03183-2. [DOI] [PMC free article] [PubMed] [Google Scholar]
37.Tasaka G, et al. The Temporal Association Cortex Plays a Key Role in Auditory-Driven Maternal Plasticity. Neuron. 2020;107:566–579.:e7. doi: 10.1016/j.neuron.2020.05.004. [ This study identifies the temporal association cortex as a key cortical hub for processing pup vocalisations and shows that maternal experience enhances the discriminability of pup calls in this area ] [DOI] [PMC free article] [PubMed] [Google Scholar]
38.Valtcheva S, et al. Neural circuitry for maternal oxytocin release induced by infant cries. Nature. 2023;621:788–795. doi: 10.1038/s41586-023-06540-4. [ Using in vivo recordings, this elegant study discovers that oxytocin neurons respond to pup vocalisations and that their activation is gated by disinhibition from the posterior intralaminar thalamus ] [DOI] [PMC free article] [PubMed] [Google Scholar]
39.LeMessurier AM, Agha AA, Stephens J, Froemke RC. Auditory corticofugal circuits drive vocalization-guided maternal behavior. BioRxiv Prepr Serv Biol. 2023:2023.09.13.557594. doi: 10.1101/2023.09.13.557594. [DOI] [Google Scholar]
40.Matsuo T, et al. Genetic dissection of pheromone processing reveals main olfactory system-mediated social behaviors in mice. Proc Natl Acad Sci U S A. 2015;112:E311–320. doi: 10.1073/pnas.1416723112. [DOI] [PMC free article] [PubMed] [Google Scholar]
41.Wang Z, Storm DR. Maternal Behavior is Impaired in Female Mice Lacking Type 3 Adenylyl Cyclase. Neuropsychopharmacology. 2011;36:772–781. doi: 10.1038/npp.2010.211. [DOI] [PMC free article] [PubMed] [Google Scholar]
42.Zarrow M, Gandelman R, Denenberg VH. Lack of nest building and maternal behavior in the mouse following olfactory bulb removal. Horm Behav. 1971;2:227–238. [Google Scholar]
43.Gandelman R, Zarrow MX, Denenberg VH, Myers M. Olfactory Bulb Removal Eliminates Maternal Behavior in the Mouse. Science. 1971;171:210–211. doi: 10.1126/science.171.3967.210. [DOI] [PubMed] [Google Scholar]
44.Canteras NS, Simerly RB, Swanson LW. Organization of projections from the medial nucleus of the amygdala: a PHAL study in the rat. J Comp Neurol. 1995;360:213–245. doi: 10.1002/cne.903600203. [DOI] [PubMed] [Google Scholar]
45.Raam T, Hong W. Organization of neural circuits underlying social behavior: A consideration of the medial amygdala. Curr Opin Neurobiol. 2021;68:124–136. doi: 10.1016/j.conb.2021.02.008. [DOI] [PMC free article] [PubMed] [Google Scholar]
46.Chen PB, et al. Sexually Dimorphic Control of Parenting Behavior by the Medial Amygdala. Cell. 2019;176:1206–1221.:e18. doi: 10.1016/j.cell.2019.01.024. [DOI] [PMC free article] [PubMed] [Google Scholar]
47.Kohl J, et al. Functional circuit architecture underlying parental behaviour. Nature. 2018;556:326–331. doi: 10.1038/s41586-018-0027-0. [DOI] [PMC free article] [PubMed] [Google Scholar]
48.Kohl J, Dulac C. Neural control of parental behaviors. Curr Opin Neurobiol. 2018;49:116–122. doi: 10.1016/j.conb.2018.02.002. [DOI] [PMC free article] [PubMed] [Google Scholar]
49.Keller M, Perrin G, Meurisse M, Ferreira G, Lévy F. Cortical and medial amygdala are both involved in the formation of olfactory offspring memory in sheep. Eur J Neurosci. 2004;20:3433–3441. doi: 10.1111/j.1460-9568.2004.03812.x. [DOI] [PubMed] [Google Scholar]
50.Nowak R, Keller M, Lévy F. Mother–Young Relationships in Sheep: A Model for a Multidisciplinary Approach of the Study of Attachment in Mammals. J Neuroendocrinol. 2011;23:1042–1053. doi: 10.1111/j.1365-2826.2011.02205.x. [DOI] [PubMed] [Google Scholar]
51.Nowlan AC, et al. Multisensory integration of social signals by a pathway from the basal amygdala to the auditory cortex in maternal mice. Curr Biol. 2025;35:36–49.:e4. doi: 10.1016/j.cub.2024.10.078. [DOI] [PMC free article] [PubMed] [Google Scholar]
52.Tachikawa KS, Yoshihara Y, Kuroda KO. Behavioral Transition from Attack to Parenting in Male Mice: A Crucial Role of the Vomeronasal System. J Neurosci. 2013;33:5120–5126. doi: 10.1523/JNEUROSCI.2364-12.2013. [DOI] [PMC free article] [PubMed] [Google Scholar]
53.Wu Z, Autry AE, Bergan JF, Watabe-Uchida M, Dulac CG. Galanin neurons in the medial preoptic area govern parental behaviour. Nature. 2014;509:325–330. doi: 10.1038/nature13307. [DOI] [PMC free article] [PubMed] [Google Scholar]
54.Prakash N, Abu Irqeba A, Corbin JG. Development and function of the medial amygdala. Trends Neurosci. 2025;48:22–32. doi: 10.1016/j.tins.2024.11.004. [DOI] [PMC free article] [PubMed] [Google Scholar]
55.Keshavarzi S, Sullivan RKP, Ianno DJ, Sah P. Functional properties and projections of neurons in the medial amygdala. J Neurosci Off J Soc Neurosci. 2014;34:8699–8715. doi: 10.1523/JNEUROSCI.1176-14.2014. [DOI] [PMC free article] [PubMed] [Google Scholar]
56.Sheehan T, Paul M, Amaral E, Numan MJ, Numan M. Evidence that the medial amygdala projects to the anterior/ventromedial hypothalamic nuclei to inhibit maternal behavior in rats. Neuroscience. 2001;106:341–356. doi: 10.1016/s0306-4522(01)00286-x. [DOI] [PubMed] [Google Scholar]
57.Xie Z, et al. Mechanically evoked defensive attack is controlled by GABAergic neurons in the anterior hypothalamic nucleus. Nat Neurosci. 2022;25:72–85. doi: 10.1038/s41593-021-00985-4. [DOI] [PubMed] [Google Scholar]
58.Wang L, Chen IZ, Lin D. Collateral Pathways from the Ventromedial Hypothalamus Mediate Defensive Behaviors. Neuron. 2015;85:1344–1358. doi: 10.1016/j.neuron.2014.12.025. [DOI] [PMC free article] [PubMed] [Google Scholar]
59.Lischinsky JE, et al. Transcriptionally defined amygdala subpopulations play distinct roles in innate social behaviors. Nat Neurosci. 2023;26:2131–2146. doi: 10.1038/s41593-023-01475-5. [DOI] [PMC free article] [PubMed] [Google Scholar]
60.Kenyon P, Cronin P, Keeble S. Disruption of maternal retrieving by perioral anesthesia. Physiol Behav. 1981;27:313–321. doi: 10.1016/0031-9384(81)90274-2. [DOI] [PubMed] [Google Scholar]
61.Kenyon P, Cronin P, Keeble S. Role of the infraorbital nerve in retrieving behavior in lactating rats. Behav Neurosci. 1983;97:255–269. doi: 10.1037//0735-7044.97.2.255. [DOI] [PubMed] [Google Scholar]
62.Salay LD, Tsao DY, Anderson DJ. A circuit that integrates drive state and social contact to gate mating. Nature. 2025:1–10. doi: 10.1038/s41586-025-09327-x. [DOI] [PMC free article] [PubMed] [Google Scholar]
63.Cservenák M, et al. Tuberoinfundibular Peptide of 39 Residues Is Activated during Lactation and Participates in the Suckling-Induced Prolactin Release in Rat. Endocrinology. 2010;151:5830–5840. doi: 10.1210/en.2010-0767. [DOI] [PMC free article] [PubMed] [Google Scholar]
64.Cservenák M, et al. Thalamic neuropeptide mediating the effects of nursing on lactation and maternal motivation. Psychoneuroendocrinology. 2013;38:3070–3084. doi: 10.1016/j.psyneuen.2013.09.004. [DOI] [PMC free article] [PubMed] [Google Scholar]
65.Cservenák M, et al. A Thalamo-Hypothalamic Pathway That Activates Oxytocin Neurons in Social Contexts in Female Rats. Endocrinology. 2017;158:335–348. doi: 10.1210/en.2016-1645. [DOI] [PMC free article] [PubMed] [Google Scholar]
66.Scott N, Prigge M, Yizhar O, Kimchi T. A sexually dimorphic hypothalamic circuit controls maternal care and oxytocin secretion. Nature. 2015;525:519–522. doi: 10.1038/nature15378. [DOI] [PubMed] [Google Scholar]
67.Cservenák M, et al. Maternally involved galanin neurons in the preoptic area of the rat. Brain Struct Funct. 2017;222:781–798. doi: 10.1007/s00429-016-1246-5. [DOI] [PMC free article] [PubMed] [Google Scholar]
68.Rosselet C, Zennou-Azogui Y, Xerri C. Nursing-Induced Somatosensory Cortex Plasticity: Temporally Decoupled Changes in Neuronal Receptive Field Properties Are Accompanied by Modifications in Activity-Dependent Protein Expression. J Neurosci. 2006;26:10667–10676. doi: 10.1523/JNEUROSCI.3253-06.2006. [DOI] [PMC free article] [PubMed] [Google Scholar]
69.Okabe S, et al. Pup odor and ultrasonic vocalizations synergistically stimulate maternal attention in mice. Behav Neurosci. 2013;127:432–438. doi: 10.1037/a0032395. [DOI] [PubMed] [Google Scholar]
70.Lecca S, et al. A neural substrate for negative affect dictates female parental behavior. Neuron. 2023;111:1094–1103.:e8. doi: 10.1016/j.neuron.2023.01.003. [ This study demonstrates that pup vocalisations carry negative valence in virgin female mice ] [DOI] [PubMed] [Google Scholar]
71.Carta I, et al. Sex-specific hypothalamic neural projection activity drives caregiving in mice. Nat Commun. 2025;16:4116. doi: 10.1038/s41467-025-59352-7. [DOI] [PMC free article] [PubMed] [Google Scholar]
72.Gómora-Arrati P, Dominguez G, Ågmo A. GABA Receptors in the Medial Preoptic Area Modulate the Onset of Oestradiol-Induced Maternal Behaviour in Hysterectomised-Ovariectomised, Pregnant Rats. J Neuroendocrinol. 2016;28:jne.12434. doi: 10.1111/jne.12434. [DOI] [PubMed] [Google Scholar]
73.Numan M, Numan M. A lesion and neuroanatomical tract-tracing analysis of the role of the bed nucleus of the stria terminalis in retrieval behavior and other aspects of maternal responsiveness in rats. Dev Psychobiol. 1996;29:23–51. doi: 10.1002/(SICI)1098-2302(199601)29:1<23::AID-DEV2>3.0.CO;2-O. [DOI] [PubMed] [Google Scholar]
74.Perrin G, Meurisse M, Levy F. Inactivation of the medial preoptic area or the bed nucleus of the stria terminalis differentially disrupts maternal behavior in sheep. Horm Behav. 2007;52:461–473. doi: 10.1016/j.yhbeh.2007.06.010. [DOI] [PubMed] [Google Scholar]
75.Benedict J, Cudmore RH, Oden D, Spruell A, Linden DJ. The Lateral Habenula Is Necessary for Maternal Behavior in the Naturally Parturient Primiparous Mouse Dam. eneuro. 2025;12:ENEURO.0092-24.2024. doi: 10.1523/ENEURO.0092-24.2024. [DOI] [PMC free article] [PubMed] [Google Scholar]
76.Numan M. Medial preoptic area and maternal behavior in the female rat. J Comp Physiol Psychol. 1974;87:746–759. doi: 10.1037/h0036974. [DOI] [PubMed] [Google Scholar]
77.Jacobson CD, Terkel J, Gorski RA, Sawyer CH. Effects of small medial preoptic area lesions on maternal behavior: Retreiving and nest building in the rat. Brain Res. 1980;194:471–478. doi: 10.1016/0006-8993(80)91226-3. [DOI] [PubMed] [Google Scholar]
78.Tsuneoka Y, et al. Functional, anatomical, and neurochemical differentiation of medial preoptic area subregions in relation to maternal behavior in the mouse. J Comp Neurol. 2013;521:1633–1663. doi: 10.1002/cne.23251. [DOI] [PubMed] [Google Scholar]
79.Tsuneoka Y, et al. Distinct preoptic-BST nuclei dissociate paternal and infanticidal behavior in mice. EMBO J. 2015;34:2652–2670. doi: 10.15252/embj.201591942. [ This study finds that central MPOA lesions promote infanticidal behavior in fathers, while lesions of the rhomboid nucleus of the BNST inhibit infanticide in virgin males ] [DOI] [PMC free article] [PubMed] [Google Scholar]
80.Kurachi T, et al. Behavioral competition between infant care and sexual behavior in male but not female common marmosets. Neurosci Res. 2025;216:104900. doi: 10.1016/j.neures.2025.04.003. [DOI] [PubMed] [Google Scholar]
81.Bridges RS. Neuroendocrine regulation of maternal behavior. Front Neuroendocrinol. 2015;36:178–196. doi: 10.1016/j.yfrne.2014.11.007. [DOI] [PMC free article] [PubMed] [Google Scholar]
82.Fang Y-Y, Yamaguchi T, Song SC, Tritsch NX, Lin D. A Hypothalamic Midbrain Pathway Essential for Driving Maternal Behaviors. Neuron. 2018;98:192–207.:e10. doi: 10.1016/j.neuron.2018.02.019. [DOI] [PMC free article] [PubMed] [Google Scholar]
83.Xie Y, Huang L, Corona A, Pagliaro AH, Shea SD. A dopaminergic reward prediction error signal shapes maternal behavior in mice. Neuron. 2023;111:557–570.:e7. doi: 10.1016/j.neuron.2022.11.019. [ This elegant study establishes that dopaminergic VTA neurons refine stereotyped pup retrieval behaviour by a reinforcement learning mechanism ] [DOI] [PMC free article] [PubMed] [Google Scholar]
84.Moffitt JR, et al. Molecular, spatial, and functional single-cell profiling of the hypothalamic preoptic region. Science. 2018;362:eaau5324. doi: 10.1126/science.aau5324. [DOI] [PMC free article] [PubMed] [Google Scholar]
85.Wei Y-C, et al. Medial preoptic area in mice is capable of mediating sexually dimorphic behaviors regardless of gender. Nat Commun. 2018;9:279. doi: 10.1038/s41467-017-02648-0. [DOI] [PMC free article] [PubMed] [Google Scholar]
86.Zhang Z, et al. Estrogen-sensitive medial preoptic area neurons coordinate torpor in mice. Nat Commun. 2020;11:6378. doi: 10.1038/s41467-020-20050-1. [DOI] [PMC free article] [PubMed] [Google Scholar]
87.Yoshihara C, et al. Calcitonin receptor signaling in the medial preoptic area enables risk-taking maternal care. Cell Rep. 2021;35:109204. doi: 10.1016/j.celrep.2021.109204. [DOI] [PubMed] [Google Scholar]
88.Nair A, et al. The neural computation of affective internal states in the hypothalamus: A dynamical systems perspective. Neuron. 2025;113:3887–3907. doi: 10.1016/j.neuron.2025.11.003. [DOI] [PubMed] [Google Scholar]
89.Autry AE, et al. Urocortin-3 neurons in the mouse perifornical area promote infant-directed neglect and aggression. eLife. 2021;10:e64680. doi: 10.7554/eLife.64680. [DOI] [PMC free article] [PubMed] [Google Scholar]
90.Sato K, et al. Amygdalohippocampal Area Neurons That Project to the Preoptic Area Mediate Infant-Directed Attack in Male Mice. J Neurosci. 2020;40:3981–3994. doi: 10.1523/JNEUROSCI.0438-19.2020. [DOI] [PMC free article] [PubMed] [Google Scholar]
91.Mei L, et al. The neural control of infanticide and parental behaviors in male mice. 2026 doi: 10.64898/2026.03.12.711470. Preprint at. [DOI] [Google Scholar]
92.Lee A, Clancy S, Fleming AS. Mother rats bar-press for pups: effects of lesions of the mpoa and limbic sites on maternal behavior and operant responding for pup-reinforcement. Behav Brain Res. 1999;100:15–31. doi: 10.1016/s0166-4328(98)00109-0. [DOI] [PubMed] [Google Scholar]
93.Mattson BJ, Morrell JI. Preference for cocaine-versus pup-associated cues differentially activates neurons expressing either Fos or cocaine- and amphetamine-regulated transcript in lactating, maternal rodents. Neuroscience. 2005;135:315–328. doi: 10.1016/j.neuroscience.2005.06.045. [DOI] [PMC free article] [PubMed] [Google Scholar]
94.Feldman R. The adaptive human parental brain: implications for children’s social development. Trends Neurosci. 2015;38:387–399. doi: 10.1016/j.tins.2015.04.004. [DOI] [PubMed] [Google Scholar]
95.Day KR, Shea SD. Dopamine dynamics underlying maternal motivation and reward. Neurosci Res. 2025;218:104928. doi: 10.1016/j.neures.2025.104928. [DOI] [PMC free article] [PubMed] [Google Scholar]
96.Hansen S, Harthon C, Wallin E, Löfberg L, Svensson K. Mesotelencephalic dopamine system and reproductive behavior in the female rat: Effects of ventral tegmental 6-hydroxydopamine lesions on maternal and sexual responsiveness. Behav Neurosci. 1991;105:588–598. doi: 10.1037//0735-7044.105.4.588. [DOI] [PubMed] [Google Scholar]
97.Hansen S. Maternal behavior of female rats with 6-OHDA lesions in the ventral striatum: Characterization of the pup retrieval deficit. Physiol Behav. 1994;55:615–620. doi: 10.1016/0031-9384(94)90034-5. [DOI] [PubMed] [Google Scholar]
98.Numan M, Stolzenberg DS, Dellevigne AA, Correnti CM, Numan MJ. Temporary inactivation of ventral tegmental area neurons with either muscimol or baclofen reversibly disrupts maternal behavior in rats through different underlying mechanisms. Behav Neurosci. 2009;123:740–751. doi: 10.1037/a0016204. [DOI] [PubMed] [Google Scholar]
99.Keer SE, Stern JM. Dopamine Receptor Blockade in the Nucleus Accumbens Inhibits Maternal Retrieval and Licking, but Enhances Nursing Behavior in Lactating Rats. Physiol Behav. 1999;67:659–669. doi: 10.1016/s0031-9384(99)00116-x. [DOI] [PubMed] [Google Scholar]
100.Li M, Fleming AS. Differential involvement of nucleus accumbens shell and core subregions in maternal memory in postpartum female rats. Behav Neurosci. 2003;117:426–445. doi: 10.1037/0735-7044.117.3.426. [DOI] [PubMed] [Google Scholar]
101.Champagne FA, et al. Variations in Nucleus Accumbens Dopamine Associated with Individual Differences in Maternal Behavior in the Rat. J Neurosci. 2004;24:4113–4123. doi: 10.1523/JNEUROSCI.5322-03.2004. [DOI] [PMC free article] [PubMed] [Google Scholar]
102.Alsina-Llanes M, Olazábal DE. NMDA- and 6-OHDA-induced Lesions in the Nucleus Accumbens Differently Affect Maternal and Infanticidal Behavior in Pup-naïve Female and Male Mice. Neuroscience. 2024;539:35–50. doi: 10.1016/j.neuroscience.2023.12.009. [DOI] [PubMed] [Google Scholar]
103.Strathearn L, Li J, Fonagy P, Montague PR. What’s in a Smile? Maternal Brain Responses to Infant Facial Cues. Pediatrics. 2008;122:40–51. doi: 10.1542/peds.2007-1566. [DOI] [PMC free article] [PubMed] [Google Scholar]
104.Strathearn L, Fonagy P, Amico J, Montague PR. Adult Attachment Predicts Maternal Brain and Oxytocin Response to Infant Cues. Neuropsychopharmacology. 2009;34:2655–2666. doi: 10.1038/npp.2009.103. [DOI] [PMC free article] [PubMed] [Google Scholar]
105.Tasaka G-I, et al. Orbitofrontal cortex influences dopamine dynamics associated with alloparental behavioral acquisition in female mice. Sci Adv. 2025;11:eadr4620. doi: 10.1126/sciadv.adr4620. [DOI] [PMC free article] [PubMed] [Google Scholar]
106.Stempel AV. A conserved brainstem region for instinctive behaviour control: The vertebrate periaqueductal gray. Curr Opin Neurobiol. 2024;86:102878. doi: 10.1016/j.conb.2024.102878. [DOI] [PubMed] [Google Scholar]
107.Bandler R, Shipley MT. Columnar organization in the midbrain periaqueductal gray: modules for emotional expression? Trends Neurosci. 1994;17:379–389. doi: 10.1016/0166-2236(94)90047-7. [DOI] [PubMed] [Google Scholar]
108.Lonstein JS, Stern JM. Role of the Midbrain Periaqueductal Gray in Maternal Nurturance and Aggression: c-fos and Electrolytic Lesion Studies in Lactating Rats. J Neurosci. 1997;17:3364–3378. doi: 10.1523/JNEUROSCI.17-09-03364.1997. [DOI] [PMC free article] [PubMed] [Google Scholar]
109.Lonstein JS, Stern JM. Site and behavioral specificity of periaqueductal gray lesions on postpartum sexual, maternal, and aggressive behaviors in rats. Brain Res. 1998;804:21–35. doi: 10.1016/s0006-8993(98)00642-8. [DOI] [PubMed] [Google Scholar]
110.Lonstein JS, Simmons DA, Stern JM. Functions of the caudal periaqueductal gray in lactating rats: Kyphosis, lordosis, maternal aggression, and fearfulness. Behav Neurosci. 1998;112:1502–1518. doi: 10.1037//0735-7044.112.6.1502. [DOI] [PubMed] [Google Scholar]
111.Zhang G-W, et al. Medial preoptic area antagonistically mediates stress-induced anxiety and parental behavior. Nat Neurosci. 2021;24:516–528. doi: 10.1038/s41593-020-00784-3. [DOI] [PMC free article] [PubMed] [Google Scholar]
112.Zhu Z, et al. A substantia innominata-midbrain circuit controls a general aggressive response. Neuron. 2021;109:1540–1553.:e9. doi: 10.1016/j.neuron.2021.03.002. [DOI] [PubMed] [Google Scholar]
113.Numan M, Numan MJ. Preoptic-brainstem connections and maternal behavior in rats. Behav Neurosci. 1991;105:1013–1029. doi: 10.1037//0735-7044.105.6.1013. [DOI] [PubMed] [Google Scholar]
114.Cohen L, Rothschild G, Mizrahi A. Multisensory Integration of Natural Odors and Sounds in the Auditory Cortex. Neuron. 2011;72:357–369. doi: 10.1016/j.neuron.2011.08.019. [DOI] [PubMed] [Google Scholar]
115.Afonso VM, Sison M, Lovic V, Fleming AS. Medial prefrontal cortex lesions in the female rat affect sexual and maternal behavior and their sequential organization. Behav Neurosci. 2007;121:515–526. doi: 10.1037/0735-7044.121.3.515. [DOI] [PubMed] [Google Scholar]
116.Febo M, Felix-Ortiz AC, Johnson TR. Inactivation or inhibition of neuronal activity in the medial prefrontal cortex largely reduces pup retrieval and grouping in maternal rats. Brain Res. 2010;1325:77–88. doi: 10.1016/j.brainres.2010.02.027. [DOI] [PMC free article] [PubMed] [Google Scholar]
117.Wu K, et al. Experience-dependent maternal defense behavior mediated by profrontal cortical projections to the medial preoptic area in mice. Nat Commun. 2025;16:7664. doi: 10.1038/s41467-025-63062-5. [DOI] [PMC free article] [PubMed] [Google Scholar]
118.Swain JE, et al. Approaching the biology of human parental attachment: Brain imaging, oxytocin and coordinated assessments of mothers and fathers. Brain Res. 2014;1580:78–101. doi: 10.1016/j.brainres.2014.03.007. [DOI] [PMC free article] [PubMed] [Google Scholar]
119.Musser ED, Kaiser-Laurent H, Ablow JC. The neural correlates of maternal sensitivity: An fMRI study. Dev Cogn Neurosci. 2012;2:428–436. doi: 10.1016/j.dcn.2012.04.003. [DOI] [PMC free article] [PubMed] [Google Scholar]
120.Gandelman R. The ontogeny of maternal responsiveness in female Rockland-Swiss albino mice. Horm Behav. 1973;4:257–268. doi: 10.1016/0018-506x(73)90010-x. [DOI] [PubMed] [Google Scholar]
121.Gandelman R. The development of cannibalism in male Rockland-Swiss mice and the influence of olfactory bulb removal. Dev Psychobiol. 1973;6:159–164. doi: 10.1002/dev.420060210. [DOI] [PubMed] [Google Scholar]
122.Rosenblatt JS. Nonhormonal Basis of Maternal Behavior in the Rat. Science. 1967;156:1512–1514. doi: 10.1126/science.156.3781.1512. [DOI] [PubMed] [Google Scholar]
123.Husain BFA, Kohl J. Plasticity of the parental brain. Curr Opin Neurobiol. 2025;95:103129. doi: 10.1016/j.conb.2025.103129. [DOI] [PubMed] [Google Scholar]
124.Servin-Barthet C, et al. The transition to motherhood: linking hormones, brain and behaviour. Nat Rev Neurosci. 2023;24:605–619. doi: 10.1038/s41583-023-00733-6. [DOI] [PubMed] [Google Scholar]
125.Hoekzema E, et al. Pregnancy leads to long-lasting changes in human brain structure. Nat Neurosci. 2017;20:287–296. doi: 10.1038/nn.4458. [DOI] [PubMed] [Google Scholar]
126.Kenkel WM, Perkeybile AM, Carter CS. The neurobiological causes and effects of alloparenting. Dev Neurobiol. 2017;77:214–232. doi: 10.1002/dneu.22465. [DOI] [PMC free article] [PubMed] [Google Scholar]
127.Feldman R, Braun K, Champagne FA. The neural mechanisms and consequences of paternal caregiving. Nat Rev Neurosci. 2019;20:205–224. doi: 10.1038/s41583-019-0124-6. [DOI] [PubMed] [Google Scholar]
128.Bridges RS, Zarrow MX, Goldman BD, Denenberg VH. A developmental study of maternal responsiveness in the rat. Physiol Behav. 1974;12:149–151. doi: 10.1016/0031-9384(74)90082-1. [DOI] [PubMed] [Google Scholar]
129.Mayer AD, Rosenblatt JS. Ontogeny of maternal behavior in the laboratory rat: Early origins in 18-to 27-day-old young. Dev Psychobiol. 1979;12:407–424. doi: 10.1002/dev.420120502. [DOI] [PubMed] [Google Scholar]
130.Roberts RL, Miller AK, Taymans SE, Carter CS. Role of social and endocrine factors in alloparental behavior of prairie voles (Microtus ochrogaster) Can J Zool. 1998;76:1862–1868. [Google Scholar]
131.Olazábal DE, Young LJ. Variability in “spontaneous” maternal behavior is associated with anxiety-like behavior and affiliation in Naïve juvenile and adult female prairie voles (Microtus ochrogaster) Dev Psychobiol. 2005;47:166–178. doi: 10.1002/dev.20077. [DOI] [PubMed] [Google Scholar]
132.Kaplan HS, et al. Sensory input, sex and function shape hypothalamic cell type development. Nature. 2025;647:157–168. doi: 10.1038/s41586-025-08603-0. [DOI] [PMC free article] [PubMed] [Google Scholar]
133.Kercmar J, Snoj T, Tobet SA, Majdic G. Gonadectomy prior to puberty decreases normal parental behavior in adult mice. Horm Behav. 2014;66:667–673. doi: 10.1016/j.yhbeh.2014.09.007. [DOI] [PMC free article] [PubMed] [Google Scholar]
134.Fukui K, Uki H, Minami M, Amano T. Effect of gonadal steroid hormone levels during pubertal development on social behavior of adult mice toward pups and synaptic transmission in the rhomboid nucleus of the bed nucleus of the stria terminalis. Neurosci Lett. 2019;708:134357. doi: 10.1016/j.neulet.2019.134357. [DOI] [PubMed] [Google Scholar]
135.Weissner B, Sheard N. Maternal behaviour in the rat. Endocrinology. 1933;17:590. [Google Scholar]
136.Leblond CP. Extra-Hormonal Factors in Maternal Behavior. Exp Biol Med. 1938;38:66–70. [Google Scholar]
137.Alsina-Llanes M, Lenzi B, Olazábal DE. Repeated non-contact exposure to pups inhibits infanticidal and facilitates paternal behavior in virgin adult male mice (C57BL6) Physiol Behav. 2025;295:114885. doi: 10.1016/j.physbeh.2025.114885. [DOI] [PubMed] [Google Scholar]
138.Cai W, et al. Involvement of the dopamine system in paternal behavior induced by repeated pup exposure in virgin male ICR mice. Behav Brain Res. 2021;415:113519. doi: 10.1016/j.bbr.2021.113519. [DOI] [PubMed] [Google Scholar]
139.Stolzenberg DS, Rissman EF. Oestrogen-Independent, Experience-Induced Maternal Behaviour in Female Mice: Oestradiol-independent maternal behaviour. J Neuroendocrinol. 2011;23:345–354. doi: 10.1111/j.1365-2826.2011.02112.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
140.Noirot E. Changes in responsiveness to young in the adult mouse. I. The problematical effect of hormones. Anim Behav. 1964;12:52–58. [Google Scholar]
141.Noirot E. Changes in responsiveness to young in the adult mouse. IV. The effect of an initial contact with a strong stimulus. Anim Behav. 1964;12:442–445. [Google Scholar]
142.Cohen J, Bridges RS. Retention of maternal behavior in nulliparous and primiparous rats: Effects of duration of previous maternal experience. J Comp Physiol Psychol. 1981;95:450–459. [Google Scholar]
143.Bridges RS, Scanlan VF. Maternal memory in adult, nulliparous rats: Effects of testing interval on the retention of maternal behavior. Dev Psychobiol. 2005;46:13–18. doi: 10.1002/dev.20038. [DOI] [PubMed] [Google Scholar]
144.Okabe S, et al. Pup exposure facilitates retrieving behavior via the oxytocin neural system in female mice. Psychoneuroendocrinology. 2017;79:20–30. doi: 10.1016/j.psyneuen.2017.01.036. [DOI] [PubMed] [Google Scholar]
145.Bair-Marshall C, Cassel N, Agha A, Bkhiet M, Froemke R. Neural circuit plasticity transforms infant neglect into maternal care. 2025 doi: 10.21203/rs.3.rs-5983736/v1. Preprint at. [DOI] [Google Scholar]
146.Swart JM, Grattan DR, Ladyman SR, Brown RSE. Pups and prolactin are rewarding to virgin female and pregnant mice. J Neuroendocrinol. 2023;35:e13232. doi: 10.1111/jne.13232. [DOI] [PubMed] [Google Scholar]
147.Salais-López H, et al. Maternal Motivation: Exploring the Roles of Prolactin and Pup Stimuli. Neuroendocrinology. 2021;111:805–830. doi: 10.1159/000510038. [DOI] [PubMed] [Google Scholar]
148.Quinones-Laracuente K, et al. A collicular-hypothalamic pathway for social visual awareness. BioRxiv Prepr Serv Biol. 2026:2026.04.17.719268. doi: 10.64898/2026.04.17.719268. [DOI] [Google Scholar]
149.Rawn SM, et al. Pregnancy Hyperglycemia in Prolactin Receptor Mutant, but Not Prolactin Mutant, Mice and Feeding-Responsive Regulation of Placental Lactogen Genes Implies Placental Control of Maternal Glucose Homeostasis. Biol Reprod. 2015;93(75):1–12. doi: 10.1095/biolreprod.115.132431. [DOI] [PMC free article] [PubMed] [Google Scholar]
150.Soares MJ. The prolactin and growth hormone families: Pregnancy-specific hormones/cytokines at the maternal-fetal interface. Reprod Biol Endocrinol. 2004;2:51. doi: 10.1186/1477-7827-2-51. [DOI] [PMC free article] [PubMed] [Google Scholar]
151.Brown RSE, et al. Prolactin action in the medial preoptic area is necessary for postpartum maternal nursing behavior. Proc Natl Acad Sci. 2017;114:10779–10784. doi: 10.1073/pnas.1708025114. [ This study establishes a critical role for prolactin / placental lactogen action in the MPOA for maternal caregiving behaviour in mice ] [DOI] [PMC free article] [PubMed] [Google Scholar]
152.Georgescu T, Swart JM, Grattan DR, Brown RSE. The Prolactin Family of Hormones as Regulators of Maternal Mood and Behavior. Front Glob Womens Health. 2021;2:767467. doi: 10.3389/fgwh.2021.767467. [DOI] [PMC free article] [PubMed] [Google Scholar]
153.Smiley KO, Brown RSE, Grattan DR. Prolactin Action Is Necessary for Parental Behavior in Male Mice. J Neurosci. 2022;42:8308–8327. doi: 10.1523/JNEUROSCI.0558-22.2022. [ This study finds that prolactin action in the forebrain is critical for the establishment of paternal behaviour in male mice ] [DOI] [PMC free article] [PubMed] [Google Scholar]
154.Francis K, Meddle SL, Bishop VR, Russell JA. Progesterone receptor expression in the pregnant and parturient rat hypothalamus and brainstem. Brain Res. 2002;927:18–26. doi: 10.1016/s0006-8993(01)03318-2. [DOI] [PubMed] [Google Scholar]
155.Doerr HK, Siegel HI, Rosenblatt JS. Effects of progesterone withdrawal and estrogen on maternal behavior in nulliparous rats. Behav Neural Biol. 1981;32:35–44. doi: 10.1016/s0163-1047(81)90242-9. [DOI] [PubMed] [Google Scholar]
156.Murakami G. Distinct Effects of Estrogen on Mouse Maternal Behavior: The Contribution of Estrogen Synthesis in the Brain. PLOS ONE. 2016;11:e0150728. doi: 10.1371/journal.pone.0150728. [DOI] [PMC free article] [PubMed] [Google Scholar]
157.Rosenblatt JS, Wagner CK, Morrell JI. Hormonal priming and triggering of maternal behavior in the rat with special reference to the relations between estrogen receptor binding and ER mRNA in specific brain regions. Psychoneuroendocrinology. 1994;19:543–552. doi: 10.1016/0306-4530(94)90039-6. [DOI] [PubMed] [Google Scholar]
158.Glynn LM, Davis EP, Sandman CA, Goldberg WA. Gestational hormone profiles predict human maternal behavior at 1-year postpartum. Horm Behav. 2016;85:19–25. doi: 10.1016/j.yhbeh.2016.07.002. [DOI] [PMC free article] [PubMed] [Google Scholar]
159.Ribeiro AC, et al. siRNA silencing of estrogen receptor-α expression specifically in medial preoptic area neurons abolishes maternal care in female mice. Proc Natl Acad Sci. 2012;109:16324–16329. doi: 10.1073/pnas.1214094109. [DOI] [PMC free article] [PubMed] [Google Scholar]
160.Ogawa S, et al. Roles of Estrogen Receptor-α Gene Expression in Reproduction-Related Behaviors in Female Mice. Endocrinology. 1998;139:5070–5081. doi: 10.1210/endo.139.12.6357. [DOI] [PubMed] [Google Scholar]
161.Catanese MC, Vandenberg LN. Bisphenol S (BPS) alters maternal behavior and brain in mice exposed during pregnancy/lactation and their daughters. Endocrinology. 2016:en.2016-1723. doi: 10.1210/en.2016-1723. [DOI] [PMC free article] [PubMed] [Google Scholar]
162.Champagne FA, et al. Maternal Care Associated with Methylation of the Estrogen Receptor-α1b Promoter and Estrogen Receptor-α Expression in the Medial Preoptic Area of Female Offspring. Endocrinology. 2006;147:2909–2915. doi: 10.1210/en.2005-1119. [DOI] [PubMed] [Google Scholar]
163.Champagne FA, Weaver ICG, Diorio J, Sharma S, Meaney MJ. Natural Variations in Maternal Care Are Associated with Estrogen Receptor α Expression and Estrogen Sensitivity in the Medial Preoptic Area. Endocrinology. 2003;144:4720–4724. doi: 10.1210/en.2003-0564. [DOI] [PubMed] [Google Scholar]
164.Topilko T, et al. Edinger-Westphal peptidergic neurons enable maternal preparatory nesting. Neuron. 2022;110:1385–1399.:e8. doi: 10.1016/j.neuron.2022.01.012. [DOI] [PMC free article] [PubMed] [Google Scholar]
165.Brunton PJ, Russell JA. The expectant brain: adapting for motherhood. Nat Rev Neurosci. 2008;9:11–25. doi: 10.1038/nrn2280. [DOI] [PubMed] [Google Scholar]
166.Neumann I, Douglas AJ, Pittman QJ, Russell JA, Landgraf R. Oxytocin Released within the Supraoptic Nucleus of the Rat Brain by Positive Feedback Action is Involved in Parturition-Related Events. J Neuroendocrinol. 1996;8:227–233. doi: 10.1046/j.1365-2826.1996.04557.x. [DOI] [PubMed] [Google Scholar]
167.Jurek B, Neumann ID. The Oxytocin Receptor: From Intracellular Signaling to Behavior. Physiol Rev. 2018;98:1805–1908. doi: 10.1152/physrev.00031.2017. [DOI] [PubMed] [Google Scholar]
168.Menon R, Neumann ID. Detection, processing and reinforcement of social cues: regulation by the oxytocin system. Nat Rev Neurosci. 2023;24:761–777. doi: 10.1038/s41583-023-00759-w. [DOI] [PubMed] [Google Scholar]
169.Meddle SL, Bishop VR, Gkoumassi E, Van Leeuwen FW, Douglas AJ. Dynamic Changes in Oxytocin Receptor Expression and Activation at Parturition in the Rat Brain. Endocrinology. 2007;148:5095–5104. doi: 10.1210/en.2007-0615. [DOI] [PubMed] [Google Scholar]
170.Pedersen CA, Prange AJ. Induction of maternal behavior in virgin rats after intracerebroventricular administration of oxytocin. Proc Natl Acad Sci. 1979;76:6661–6665. doi: 10.1073/pnas.76.12.6661. [DOI] [PMC free article] [PubMed] [Google Scholar]
171.Pedersen CA, Caldwell JD, Johnson MF, Fort SA, Prange AJ. Oxytocin antiserum delays onset of ovarian steroid-induced maternal behavior. Neuropeptides. 1985;6:175–182. doi: 10.1016/0143-4179(85)90108-8. [DOI] [PubMed] [Google Scholar]
172.Kendrick KM, Keverne EB, Baldwin BA. Intracerebroventricular Oxytocin Stimulates Maternal Behaviour in the Sheep. Neuroendocrinology. 1987;46:56–61. doi: 10.1159/000124796. [DOI] [PubMed] [Google Scholar]
173.Fahrbach SE, Morrell JI, Pfaff DW. Possible Role for Endogenous Oxytocin in Estrogen-Facilitated Maternal Behavior in Rats. Neuroendocrinology. 1985;40:526–532. doi: 10.1159/000124125. [DOI] [PubMed] [Google Scholar]
174.Boccia ML, Goursaud A-PS, Bachevalier J, Anderson KD, Pedersen CA. Peripherally administered non-peptide oxytocin antagonist, L368,899, accumulates in limbic brain areas: A new pharmacological tool for the study of social motivation in non-human primates. Horm Behav. 2007;52:344–351. doi: 10.1016/j.yhbeh.2007.05.009. [DOI] [PMC free article] [PubMed] [Google Scholar]
175.Nishimori K, et al. Oxytocin is required for nursing but is not essential for parturition or reproductive behavior. Proc Natl Acad Sci. 1996;93:11699–11704. doi: 10.1073/pnas.93.21.11699. [DOI] [PMC free article] [PubMed] [Google Scholar]
176.Takayanagi Y, et al. Pervasive social deficits, but normal parturition, in oxytocin receptor-deficient mice. Proc Natl Acad Sci. 2005;102:16096–16101. doi: 10.1073/pnas.0505312102. [DOI] [PMC free article] [PubMed] [Google Scholar]
177.Pedersen CA, Vadlamudi SV, Boccia ML, Amico JA. Maternal behavior deficits in nulliparous oxytocin knockout mice. Genes Brain Behav. 2006;5:274–281. doi: 10.1111/j.1601-183X.2005.00162.x. [DOI] [PubMed] [Google Scholar]
178.Rich ME, deCárdenas EJ, Lee H-J, Caldwell HK. Impairments in the Initiation of Maternal Behavior in Oxytocin Receptor Knockout Mice. PLoS ONE. 2014;9:e98839. doi: 10.1371/journal.pone.0098839. [DOI] [PMC free article] [PubMed] [Google Scholar]
179.Tsuneoka Y, et al. Oxytocin Facilitates Allomaternal Behavior under Stress in Laboratory Mice. eneuro. 2022;9:ENEURO.0405-21.2022. doi: 10.1523/ENEURO.0405-21.2022. [DOI] [PMC free article] [PubMed] [Google Scholar]
180.Labriola J. Effects of Caesarean Delivery upon Maternal Behavior in Rats. Exp Biol Med. 1953;83:556–557. doi: 10.3181/00379727-83-20416. [DOI] [PubMed] [Google Scholar]
181.Moltz H, Robbins D, Parks M. Caesarean delivery and maternal behavior of primiparous and multiparous rats. J Comp Physiol Psychol. 1966;61:455–460. doi: 10.1037/h0023263. [DOI] [PubMed] [Google Scholar]
182.Li T, et al. Intranasal Oxytocin Restores Maternal Behavior and Oxytocin Neuronal Activity in the Supraoptic Nucleus in Rat Dams with Cesarean Delivery. Neuroscience. 2021;468:235–246. doi: 10.1016/j.neuroscience.2021.06.020. [DOI] [PubMed] [Google Scholar]
183.Arnoux I, et al. Astrocytes control oxytocin-based maternal behavior via connexin 30. 2026 doi: 10.64898/2026.01.09.698569. Preprint at. [DOI] [Google Scholar]
184.McNeilly AS, Robinson IC, Houston MJ, Howie PW. Release of oxytocin and prolactin in response to suckling. Br Med J. 1983;286:257–259. doi: 10.1136/bmj.286.6361.257. [DOI] [PMC free article] [PubMed] [Google Scholar]
185.Lucas BK, Ormandy CJ, Binart N, Bridges RS, Kelly PA. Null Mutation of the Prolactin Receptor Gene Produces a Defect in Maternal Behavior. Endocrinology. 1998;139 doi: 10.1210/endo.139.10.6243. [DOI] [PubMed] [Google Scholar]
186.Yamaguchi T, Lin D. The hormonal and neural control of maternal aggression. Curr Opin Neurobiol. 2026;98:103194. doi: 10.1016/j.conb.2026.103194. [DOI] [PMC free article] [PubMed] [Google Scholar]
187.Stagkourakis S, et al. Maternal aggression driven by the transient mobilisation of a dormant hormone-sensitive circuit. Nat Commun. 2025;16:8553. doi: 10.1038/s41467-025-64043-4. [DOI] [PMC free article] [PubMed] [Google Scholar]
188.Yamaguchi T, et al. The neural mechanisms supporting the rise and fall of maternal aggression. Nature. 2026:1–11. doi: 10.1038/s41586-026-10354-5. [DOI] [PMC free article] [PubMed] [Google Scholar]
189.Georgescu T, Khant Aung Z, Grattan DR, Brown RSE. Prolactin-mediated restraint of maternal aggression in lactation. Proc Natl Acad Sci. 2022;119:e2116972119. doi: 10.1073/pnas.2116972119. [DOI] [PMC free article] [PubMed] [Google Scholar]
190.Shahrokh DK, Zhang T-Y, Diorio J, Gratton A, Meaney MJ. Oxytocin-Dopamine Interactions Mediate Variations in Maternal Behavior in the Rat. Endocrinology. 2010;151:2276–2286. doi: 10.1210/en.2009-1271. [DOI] [PMC free article] [PubMed] [Google Scholar]
191.Bosch OJ, Neumann ID. Brain vasopressin is an important regulator of maternal behavior independent of dams’ trait anxiety. Proc Natl Acad Sci. 2008;105:17139–17144. doi: 10.1073/pnas.0807412105. [DOI] [PMC free article] [PubMed] [Google Scholar]
192.Bosch OJ, Pförtsch J, Beiderbeck DI, Landgraf R, Neumann ID. Maternal Behaviour is Associated with Vasopressin Release in the Medial Preoptic Area and Bed Nucleus of the Stria Terminalis in the Rat. J Neuroendocrinol. 2010;22:420–429. doi: 10.1111/j.1365-2826.2010.01984.x. [DOI] [PubMed] [Google Scholar]
193.Nephew B, Bridges R. Central actions of arginine vasopressin and a V1a receptor antagonist on maternal aggression, maternal behavior, and grooming in lactating rats. Pharmacol Biochem Behav. 2008;91:77–83. doi: 10.1016/j.pbb.2008.06.013. [DOI] [PMC free article] [PubMed] [Google Scholar]
194.Caughey SD, et al. Changes in the Intensity of Maternal Aggression and Central Oxytocin and Vasopressin V1a Receptors Across the Peripartum Period in the Rat. J Neuroendocrinol. 2011;23:1113–1124. doi: 10.1111/j.1365-2826.2011.02224.x. [DOI] [PubMed] [Google Scholar]
195.Bayerl DS, Klampfl SM, Bosch OJ. Central V1b Receptor Antagonism in Lactating Rats: Impairment of Maternal Care But Not of Maternal Aggression. J Neuroendocrinol. 2014;26:918–926. doi: 10.1111/jne.12226. [DOI] [PubMed] [Google Scholar]
196.Aspesi D, et al. The deletion of the arginine vasopressin 1a receptor impairs sexual and maternal behavior. Front Endocrinol. 2025;16:1649706. doi: 10.3389/fendo.2025.1649706. [DOI] [PMC free article] [PubMed] [Google Scholar]
197.Dvorkin R, Shea SD. Precise and Pervasive Phasic Bursting in Locus Coeruleus during Maternal Behavior in Mice. J Neurosci. 2022;42:2986–2999. doi: 10.1523/JNEUROSCI.0938-21.2022. [DOI] [PMC free article] [PubMed] [Google Scholar]
198.Corona A, Choe J, Muñoz-Castañeda R, Osten P, Shea SD. A circuit from the locus coeruleus to the anterior cingulate cortex modulates offspring interactions in mice. Cell Rep. 2023;42:112771. doi: 10.1016/j.celrep.2023.112771. [DOI] [PMC free article] [PubMed] [Google Scholar]
199.Holschbach MA, Vitale EM, Lonstein JS. Serotonin-specific lesions of the dorsal raphe disrupt maternal aggression and caregiving in postpartum rats. Behav Brain Res. 2018;348:53–64. doi: 10.1016/j.bbr.2018.04.008. [DOI] [PMC free article] [PubMed] [Google Scholar]
200.Muzerelle A, et al. Dorsal raphe serotonin neurotransmission is required for the expression of nursing behavior and for pup survival. Sci Rep. 2021;11:6004. doi: 10.1038/s41598-021-84368-6. [DOI] [PMC free article] [PubMed] [Google Scholar]
201.Gao J, Wu R, Davis C, Li M. Activation of 5-HT2A receptor disrupts rat maternal behavior. Neuropharmacology. 2018;128:96–105. doi: 10.1016/j.neuropharm.2017.09.037. [DOI] [PMC free article] [PubMed] [Google Scholar]
202.Hammer RP, Jr, Mateo AR, Bridges RS. Hormonal Regulation of Medial Preoptic μ-Opiate Receptor Density before and after Parturition. Neuroendocrinology. 1992;56:38–45. doi: 10.1159/000126206. [DOI] [PubMed] [Google Scholar]
203.Mann PE, Pasternak GW, Bridges RS. Mu1 opioid receptor involvement in maternal behavior. Physiol Behav. 1990;47:133–138. doi: 10.1016/0031-9384(90)90051-5. [DOI] [PubMed] [Google Scholar]
204.Byrnes EM, Rigero BA, Bridges RS. Opioid receptor antagonism during early lactation results in the increased duration of nursing bouts. Physiol Behav. 2000;70:211–216. doi: 10.1016/s0031-9384(00)00231-6. [DOI] [PubMed] [Google Scholar]
205.Yip SH, et al. Elevated Prolactin during Pregnancy Drives a Phenotypic Switch in Mouse Hypothalamic Dopaminergic Neurons. Cell Rep. 2019;26:1787–1799.:e5. doi: 10.1016/j.celrep.2019.01.067. [DOI] [PubMed] [Google Scholar]
206.Stolzenberg DS, Mayer HS. Experience-dependent mechanisms in the regulation of parental care. Front Neuroendocrinol. 2019;54:100745. doi: 10.1016/j.yfrne.2019.04.002. [DOI] [PMC free article] [PubMed] [Google Scholar]
207.Nephew BC, Bridges RS, Lovelock DF, Byrnes EM. Enhanced maternal aggression and associated changes in neuropeptide gene expression in multiparous rats. Behav Neurosci. 2009;123:949–957. doi: 10.1037/a0016734. [DOI] [PMC free article] [PubMed] [Google Scholar]
208.Ladyman SR, Khant Aung Z, Grattan DR. Impact of Pregnancy and Lactation on the Long-Term Regulation of Energy Balance in Female Mice. Endocrinology. 2018;159:2324–2336. doi: 10.1210/en.2018-00057. [DOI] [PubMed] [Google Scholar]
209.Chechko N, Nehls S. From Pregnancy to Postpartum: The Dynamic Reorganization of the Maternal Brain. Neurosci Insights. 2025;20:26331055251315488. doi: 10.1177/26331055251315488. [DOI] [PMC free article] [PubMed] [Google Scholar]
210.Duarte-Guterman P, et al. Cellular and molecular signatures of motherhood in the adult and ageing rat brain. Open Biol. 2023;13:230217. doi: 10.1098/rsob.230217. [DOI] [PMC free article] [PubMed] [Google Scholar]
211.De Lange A-MG, et al. Population-based neuroimaging reveals traces of childbirth in the maternal brain. Proc Natl Acad Sci. 2019;116:22341–22346. doi: 10.1073/pnas.1910666116. [DOI] [PMC free article] [PubMed] [Google Scholar]
212.Pritschet L, et al. Neuroanatomical changes observed over the course of a human pregnancy. Nat Neurosci. 2024;27:2253–2260. doi: 10.1038/s41593-024-01741-0. [DOI] [PMC free article] [PubMed] [Google Scholar]
213.Brent LJN, et al. Ecological Knowledge, Leadership, and the Evolution of Menopause in Killer Whales. Curr Biol. 2015;25:746–750. doi: 10.1016/j.cub.2015.01.037. [DOI] [PubMed] [Google Scholar]
214.Nattrass S, et al. Postreproductive killer whale grandmothers improve the survival of their grandoffspring. Proc Natl Acad Sci. 2019;116:26669–26673. doi: 10.1073/pnas.1903844116. [DOI] [PMC free article] [PubMed] [Google Scholar]
215.Lahdenperä M, Mar KU, Lummaa V. Nearby grandmother enhances calf survival and reproduction in Asian elephants. Sci Rep. 2016;6:27213. doi: 10.1038/srep27213. [DOI] [PMC free article] [PubMed] [Google Scholar]
216.Chapman SN, Lahdenperä M, Pettay JE, Lynch RF, Lummaa V. Offspring fertility and grandchild survival enhanced by maternal grandmothers in a pre-industrial human society. Sci Rep. 2021;11:3652. doi: 10.1038/s41598-021-83353-3. [DOI] [PMC free article] [PubMed] [Google Scholar]
217.Alcantara IC, et al. A hypothalamic circuit that modulates feeding and parenting behaviours. Nature. 2025;645:981–990. doi: 10.1038/s41586-025-09268-5. [DOI] [PubMed] [Google Scholar]
218.Li X-Y, et al. AGRP Neurons Project to the Medial Preoptic Area and Modulate Maternal Nest-Building. J Neurosci. 2019;39:456–471. doi: 10.1523/JNEUROSCI.0958-18.2018. [DOI] [PMC free article] [PubMed] [Google Scholar]
219.Han Y, Li X-Y, Wang S-R, Wei Y-C, Xu X-H. Presence of pups suppresses hunger-induced feeding in virgin adult mice of both sexes. Neuroscience. 2017;362:228–238. doi: 10.1016/j.neuroscience.2017.08.047. [DOI] [PubMed] [Google Scholar]
220.Muroi Y, Ishii T. Neuropeptide Y is crucial for nutritional state-dependent regulation of maternal behavior. Psychoneuroendocrinology. 2015;51:392–402. doi: 10.1016/j.psyneuen.2014.09.022. [DOI] [PubMed] [Google Scholar]
221.Zoubovsky SP, et al. Chronic psychosocial stress during pregnancy affects maternal behavior and neuroendocrine function and modulates hypothalamic CRH and nuclear steroid receptor expression. Transl Psychiatry. 2020;10:6. doi: 10.1038/s41398-020-0704-2. [DOI] [PMC free article] [PubMed] [Google Scholar]
222.Smith JW, Seckl JR, Evans AT, Costall B, Smythe JW. Gestational stress induces post-partum depression-like behaviour and alters maternal care in rats. Psychoneuroendocrinology. 2004;29:227–244. doi: 10.1016/s0306-4530(03)00025-8. [DOI] [PubMed] [Google Scholar]
223.Brummelte S, Galea LAM. Chronic corticosterone during pregnancy and postpartum affects maternal care, cell proliferation and depressive-like behavior in the dam. Horm Behav. 2010;58:769–779. doi: 10.1016/j.yhbeh.2010.07.012. [DOI] [PubMed] [Google Scholar]
224.Pedersen CA, Caldwell JD, McGuire M, Evans DL. Corticotropin-releasing hormone inhibits maternal behavior and induces pup-killing. Life Sci. 1991;48:1537–1546. doi: 10.1016/0024-3205(91)90278-j. [DOI] [PubMed] [Google Scholar]
225.Klampfl SM, et al. Maternal stress and the MPOA: Activation of CRF receptor 1 impairs maternal behavior and triggers local oxytocin release in lactating rats. Neuropharmacology. 2018;133:440–450. doi: 10.1016/j.neuropharm.2018.02.019. [DOI] [PMC free article] [PubMed] [Google Scholar]
226.Klampfl SM, et al. Brain CRF-binding protein modulates aspects of maternal behavior under stressful conditions and supports a hypo-anxious state in lactating rats. Horm Behav. 2016;84:136–144. doi: 10.1016/j.yhbeh.2016.06.009. [DOI] [PubMed] [Google Scholar]
227.Rivero-Quiles C, et al. Appetitive maternal behavior in mice is resistant to perinatal chronic variable stress and perinatal chronic corticosterone treatment. Horm Behav. 2026;177:105855. doi: 10.1016/j.yhbeh.2025.105855. [DOI] [PMC free article] [PubMed] [Google Scholar]
228.Erskine MS. Solicitation behavior in the estrous female rat: a review. Horm Behav. 1989;23:473–502. doi: 10.1016/0018-506x(89)90037-8. [DOI] [PubMed] [Google Scholar]
229.Pfaff DW, Sakuma Y. Deficit in the lordosis reflex of female rats caused by lesions in the ventromedial nucleus of the hypothalamus. J Physiol. 1979;288:203–210. [PMC free article] [PubMed] [Google Scholar]
230.Gutierrez-Castellanos N, et al. A hypothalamic node for the cyclical control of female sexual rejection. Neuron. 2024:S0896-6273(24)00774–8. doi: 10.1016/j.neuron.2024.10.026. [DOI] [PubMed] [Google Scholar]
231.Inoue S, et al. Periodic Remodeling in a Neural Circuit Governs Timing of Female Sexual Behavior. Cell. 2019;179:1393–1408.:e16. doi: 10.1016/j.cell.2019.10.025. [DOI] [PMC free article] [PubMed] [Google Scholar]
232.Yin L, et al. VMHvllCckar cells dynamically control female sexual behaviors over the reproductive cycle. Neuron. 2022;110:3000–3017.:e8. doi: 10.1016/j.neuron.2022.06.026. [DOI] [PMC free article] [PubMed] [Google Scholar]
233.Ishii KK, et al. Post-ejaculatory Inhibition of Female Sexual Drive via Heterogeneous Neuronal Ensembles in the Medial Preoptic Area. eLife. 2025;12 doi: 10.7554/eLife.91765. [DOI] [PMC free article] [PubMed] [Google Scholar]
234.Sun F, et al. Shared neural substrates of prosocial and parenting behaviours. Nature. 2026 doi: 10.1038/s41586-026-10327-8. [DOI] [PMC free article] [PubMed] [Google Scholar]
235.de Waal FBM. Putting the altruism back into altruism: the evolution of empathy. Annu Rev Psychol. 2008;59:279–300. doi: 10.1146/annurev.psych.59.103006.093625. [DOI] [PubMed] [Google Scholar]
236.Niepoth N, et al. Evolution of a novel adrenal cell type that promotes parental care. Nature. 2024;629:1082–1090. doi: 10.1038/s41586-024-07423-y. [ This study discovers that monogamous oldfield mice recently evolved a novel adrenal gland cell type which promotes caregiving behaviours by converting progesterone into 20α-hydroxyprogesterone ] [DOI] [PMC free article] [PubMed] [Google Scholar]
237.Young KA, Gobrogge KL, Liu Y, Wang Z. The neurobiology of pair bonding: insights from a socially monogamous rodent. Front Neuroendocrinol. 2011;32:53–69. doi: 10.1016/j.yfrne.2010.07.006. [DOI] [PMC free article] [PubMed] [Google Scholar]
238.Rogers FD, et al. Agouti integrates environmental cues to regulate paternal behaviour. Nature. 2026;652:694–702. doi: 10.1038/s41586-026-10123-4. [DOI] [PMC free article] [PubMed] [Google Scholar]
239.Li Y, et al. Paternal behavior is controlled by preoptic Trpc5 neurons. 2026 doi: 10.64898/2026.03.20.713203. Preprint at. [DOI] [Google Scholar]
240.Vom Saal FS, Howard LS. The Regulation of Infanticide and Parental Behavior: Implications for Reproductive Success in Male Mice. Science. 1982;215:1270–1272. doi: 10.1126/science.7058349. [DOI] [PubMed] [Google Scholar]
241.Vom Saal FS. Time-contingent change in infanticide and parental behavior induced by ejaculation in male mice. Physiol Behav. 1985;34:7–15. doi: 10.1016/0031-9384(85)90069-1. [DOI] [PubMed] [Google Scholar]
242.McCarthy MM, Vom Saal FS. Inhibition of infanticide after mating by wild male house mice. Physiol Behav. 1986;36:203–209. doi: 10.1016/0031-9384(86)90004-1. [DOI] [PubMed] [Google Scholar]
243.Perrigo G, Bryant WC, Vom Saal FS. A unique neural timing system prevents male mice from harming their own offspring. Anim Behav. 1990;39:535–539. [Google Scholar]
244.Rosenfeld CS, Johnson SA, Ellersieck MR, Roberts RM. Interactions between Parents and Parents and Pups in the Monogamous California Mouse (Peromyscus californicus) PLoS ONE. 2013;8:e75725. doi: 10.1371/journal.pone.0075725. [DOI] [PMC free article] [PubMed] [Google Scholar]
245.Valera-Marín G, et al. Raised without a father: monoparental care effects over development, sexual behavior, sexual reward, and pair bonding in prairie voles. Behav Brain Res. 2021;408:113264. doi: 10.1016/j.bbr.2021.113264. [DOI] [PMC free article] [PubMed] [Google Scholar]
246.Clutton-Brock TH, et al. Effects of Helpers on Juvenile Development and Survival in Meerkats. Science. 2001;293:2446–2449. doi: 10.1126/science.1061274. [DOI] [PubMed] [Google Scholar]
247.Ziegler TE, Prudom SL, Zahed SR. Variations in male parenting behavior and physiology in the common marmoset. Am J Hum Biol. 2009;21:739–744. doi: 10.1002/ajhb.20920. [DOI] [PMC free article] [PubMed] [Google Scholar]
248.Lukas D, Clutton-Brock T. Life histories and the evolution of cooperative breeding in mammals. Proc R Soc B Biol Sci. 2012;279:4065–4070. doi: 10.1098/rspb.2012.1433. [DOI] [PMC free article] [PubMed] [Google Scholar]
249.Russell AF, Young AJ, Spong G, Jordan NR, Clutton-Brock TH. Helpers increase the reproductive potential of offspring in cooperative meerkats. Proc R Soc B Biol Sci. 2007;274:513–520. doi: 10.1098/rspb.2006.3698. [DOI] [PMC free article] [PubMed] [Google Scholar]
250.Rosenbaum S, Gettler LT. With a little help from her friends (and family) part I: the ecology and evolution of non-maternal care in mammals. Physiol Behav. 2018;193:1–11. doi: 10.1016/j.physbeh.2017.12.025. [DOI] [PubMed] [Google Scholar]
251.Kentner AC, Abizaid A, Bielajew C. Modeling Dad: Animal models of paternal behavior. Neurosci Biobehav Rev. 2010;34:438–451. doi: 10.1016/j.neubiorev.2009.08.010. [DOI] [PubMed] [Google Scholar]
252.Gubernick DJ, Alberts JR. The biparental care system of the California mouse, Peromyscus californicus. J Comp Psychol. 1987;101:169–177. [PubMed] [Google Scholar]
253.Ahern TH, Hammock EAD, Young LJ. Parental division of labor, coordination, and the effects of family structure on parenting in monogamous prairie voles (Microtus ochrogaster) Dev Psychobiol. 2011;53:118–131. doi: 10.1002/dev.20498. [DOI] [PMC free article] [PubMed] [Google Scholar]
254.Bendesky A, et al. The genetic basis of parental care evolution in monogamous mice. Nature. 2017;544:434–439. doi: 10.1038/nature22074. [DOI] [PMC free article] [PubMed] [Google Scholar]
255.Khadraoui M, Merritt JR, Hoekstra HE, Bendesky A. Post-mating parental behavior trajectories differ across four species of deer mice. PLOS ONE. 2022;17:e0276052. doi: 10.1371/journal.pone.0276052. [DOI] [PMC free article] [PubMed] [Google Scholar]
256.Elwood RW. Changes in the responses of male and female gerbils (Meriones unguiculatus) towards test pups during the pregnancy of the female. Anim Behav. 1977;25:46–51. [Google Scholar]
257.Elwood RW, Stolzenberg DS. Flipping the parental switch: from killing to caring in male mammals. Anim Behav. 2020;165:133–142. doi: 10.1016/j.anbehav.2020.05.001. [DOI] [PMC free article] [PubMed] [Google Scholar]
258.Kennedy HF, Elwood RW. Strain differences in the inhibition of infanticide in male mice (Mus musculus) Behav Neural Biol. 1988;50:349–353. doi: 10.1016/s0163-1047(88)91066-7. [DOI] [PubMed] [Google Scholar]
259.Palanza P, Parmigiani S. Inhibition of infanticide in male Swiss mice: Behavioral polymorphism in response to multiple mediating factors. Physiol Behav. 1991;49:797–802. doi: 10.1016/0031-9384(91)90320-n. [DOI] [PubMed] [Google Scholar]
260.Lee AW, Brown RE. Medial preoptic lesions disrupt parental behavior in both male and female California mice (Peromyscus californicus) Behav Neurosci. 2002;116:968–975. doi: 10.1037//0735-7044.116.6.968. [DOI] [PubMed] [Google Scholar]
261.Lee AW, Brown RE. Comparison of medial preoptic, amygdala, and nucleus accumbens lesions on parental behavior in California mice (Peromyscus californicus) Physiol Behav. 2007;92:617–628. doi: 10.1016/j.physbeh.2007.05.008. [DOI] [PubMed] [Google Scholar]
262.Ito K, et al. Synaptic plasticity in the medial preoptic area of male mice encodes social experiences with female and regulates behavior toward young. eLife. 2025;14 [ This study proposes a two-stage model in which distinct forms of synaptic plasticity in the MPOA sequentially suppress infanticide and promote the emergence of paternal care after mating in mice ] [Google Scholar]
263.Inada K, et al. Vasopressin-to-oxytocin receptor crosstalk in the preoptic area underlying parental behaviors in male mice. Nat Commun. 2025;16:10844. doi: 10.1038/s41467-025-66908-0. [DOI] [PMC free article] [PubMed] [Google Scholar]
264.He Z, et al. Paraventricular Nucleus Oxytocin Subsystems Promote Active Paternal Behaviors in Mandarin Voles. J Neurosci Off J Soc Neurosci. 2021;41:6699–6713. doi: 10.1523/JNEUROSCI.2864-20.2021. [DOI] [PMC free article] [PubMed] [Google Scholar]
265.Trainor BC, Marler CA. Testosterone, Paternal Behavior, and Aggression in the Monogamous California Mouse (Peromyscus californicus) Horm Behav. 2001;40:32–42. doi: 10.1006/hbeh.2001.1652. [DOI] [PubMed] [Google Scholar]
266.Trainor BC, Bird IM, Alday NA, Schlinger BA, Marler CA. Variation in Aromatase Activity in the Medial Preoptic Area and Plasma Progesterone Is Associated with the Onset of Paternal Behavior. Neuroendocrinology. 2003;78:36–44. doi: 10.1159/000071704. [DOI] [PMC free article] [PubMed] [Google Scholar]
267.Gleason ED, Marler CA. Testosterone response to courtship predicts future paternal behavior in the California mouse, Peromyscus californicus. Horm Behav. 2010;57:147–154. doi: 10.1016/j.yhbeh.2009.10.006. [DOI] [PMC free article] [PubMed] [Google Scholar]
268.Wang Z, De Vries GJ. Testosterone effects on paternal behavior and vasopressin immunoreactive projections in prairie voles (Microtus ochrogaster) Brain Res. 1993;631:156–160. doi: 10.1016/0006-8993(93)91203-5. [DOI] [PubMed] [Google Scholar]
269.Lonstein, De Vries Sex Differences in the Parental Behaviour of Adult Virgin Prairie Voles: Independence From Gonadal Hormones and Vasopressin. J Neuroendocrinol. 1999;11:441–449. doi: 10.1046/j.1365-2826.1999.00361.x. [DOI] [PubMed] [Google Scholar]
270.Hume JM, Wynne-Edwards KE. Castration reduces male testosterone, estradiol, and territorial aggression, but not paternal behavior in biparental dwarf hamsters (Phodopus campbelli) Horm Behav. 2005;48:303–310. doi: 10.1016/j.yhbeh.2005.04.001. [DOI] [PubMed] [Google Scholar]
271.Saxbe DE, et al. Fathers’ decline in testosterone and synchrony with partner testosterone during pregnancy predicts greater postpartum relationship investment. Horm Behav. 2017;90:39–47. doi: 10.1016/j.yhbeh.2016.07.005. [DOI] [PubMed] [Google Scholar]
272.Meijer WM, Van IJzendoorn MH, Bakermans-Kranenburg MJ. Challenging the challenge hypothesis on testosterone in fathers: Limited meta-analytic support. Psychoneuroendocrinology. 2019;110:104435. doi: 10.1016/j.psyneuen.2019.104435. [DOI] [PubMed] [Google Scholar]
273.Gettler LT, McDade TW, Feranil AB, Kuzawa CW. Longitudinal evidence that fatherhood decreases testosterone in human males. Proc Natl Acad Sci. 2011;108:16194–16199. doi: 10.1073/pnas.1105403108. [DOI] [PMC free article] [PubMed] [Google Scholar]
274.Delahunty KM, McKay DW, Noseworthy DE, Storey AE. Prolactin responses to infant cues in men and women: Effects of parental experience and recent infant contact. Horm Behav. 2007;51:213–220. doi: 10.1016/j.yhbeh.2006.10.004. [DOI] [PubMed] [Google Scholar]
275.Gordon I, Zagoory-Sharon O, Leckman JF, Feldman R. Prolactin, Oxytocin, and the development of paternal behavior across the first six months of fatherhood. Horm Behav. 2010;58:513–518. doi: 10.1016/j.yhbeh.2010.04.007. [DOI] [PMC free article] [PubMed] [Google Scholar]
276.Feldman R, Gordon I, Schneiderman I, Weisman O, Zagoory-Sharon O. Natural variations in maternal and paternal care are associated with systematic changes in oxytocin following parent–infant contact. Psychoneuroendocrinology. 2010;35:1133–1141. doi: 10.1016/j.psyneuen.2010.01.013. [DOI] [PubMed] [Google Scholar]

[R1] 1.Stearns SC. The Evolution of Life Histories. Oxford University Press; Oxford, New York: 1992. [Google Scholar]

[R2] 2.Trivers RL. In: Foundations of Animal Behavior: Classic Papers with Commentaries. Houck LD, Drickamer LC, editors. University of Chicago Press; Chicago, IL: 1996. Parental Investment and Sexual Selection. [Google Scholar]

[R3] 3.Clutton-Brock TH. The Evolution of Parental Care. Princeton University Press; Princeton, N.J: 1991. [Google Scholar]

[R4] 4.Numan M, Insel T. The Neurobiology of Parental Behavior. Springer New York; New York, NY: 2003. [DOI] [Google Scholar]

[R5] 5.Numan M. The Parental Brain: Mechanisms, Development, and Evolution. Oxford University Press; New York: 2020. [DOI] [Google Scholar]

[R6] 6.Kaplan HS, Horvath PM, Rahman MM, Dulac C. The neurobiology of parenting and infant-evoked aggression. Physiol Rev. 2025;105:315–381. doi: 10.1152/physrev.00036.2023. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R7] 7.Maciejewski MF, Bell AM. Insights into Parental Care from Studies on Non-mammalian Vertebrates. Affect Sci. 2022;3:792–798. doi: 10.1007/s42761-022-00127-4. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R8] 8.Fischer EK, et al. The neural basis of tadpole transport in poison frogs. Proc Biol Sci. 2019;286:20191084. doi: 10.1098/rspb.2019.1084. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R9] 9.Roland AB, O’Connell LA. Poison frogs as a model system for studying the neurobiology of parental care. Curr Opin Behav Sci. 2015;6:76–81. [Google Scholar]

[R10] 10.Cunha-Saraiva F, et al. Parental care and neuropeptide dynamics in a cichlid fish Neolamprologus caudopunctatus. Horm Behav. 2019;116:104576. doi: 10.1016/j.yhbeh.2019.104576. [DOI] [PubMed] [Google Scholar]

[R11] 11.Ammari R, et al. Hormone-mediated neural remodeling orchestrates parenting onset during pregnancy. Science. 2023;382:76–81. doi: 10.1126/science.adi0576. [ This study finds that remodelling of Galanin-expressing MPOA neurons by the ovarian hormones oestradiol and progesterone is critical for the onset of parental behaviour during pregnancy in mice ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R12] 12.Cao M, et al. Integration of hunger and hormonal state gates infant-directed aggression. Nature. 2025;648:138–145. doi: 10.1038/s41586-025-09651-2. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R13] 13.Miyamichi K. Neural basis for behavioral plasticity during the parental life-stage transition in mice. Front Neural Circuits. 2024;17:1340497. doi: 10.3389/fncir.2023.1340497. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R14] 14.Mei L, Yan R, Yin L, Sullivan RM, Lin D. Antagonistic circuits mediating infanticide and maternal care in female mice. Nature. 2023;618:1006–1016. doi: 10.1038/s41586-023-06147-9. [ This study establishes that reciprocal inhibition between oestrogen receptor α (ESR1)-expressing cells in the MPOA and BNST control the balance between pup-directed care vs aggression in female mice ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R15] 15.Stolzenberg DS, Champagne FA. Hormonal and non-hormonal bases of maternal behavior: The role of experience and epigenetic mechanisms. Horm Behav. 2016;77:204–210. doi: 10.1016/j.yhbeh.2015.07.005. [DOI] [PubMed] [Google Scholar]

[R16] 16.Carcea I, et al. Oxytocin neurons enable social transmission of maternal behaviour. Nature. 2021;596:553–557. doi: 10.1038/s41586-021-03814-7. [ This study identifies oxytocin neurons as a neural substrate for the social transmission of maternal behaviour through experience-dependent cortical plasticity ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R17] 17.Inada K, et al. Plasticity of neural connections underlying oxytocin-mediated parental behaviors of male mice. Neuron. 2022;110:2009–2023.:e5. doi: 10.1016/j.neuron.2022.03.033. [ This study demonstrates that fatherhood remodels long-range synaptic inputs onto hypothalamic oxytocin neurons, enabling the transition from pup-directed aggression to paternal caregiving in male mice ] [DOI] [PubMed] [Google Scholar]

[R18] 18.Stagkourakis S, et al. A Neuro-hormonal Circuit for Paternal Behavior Controlled by a Hypothalamic Network Oscillation. Cell. 2020;182:960–975.:e15. doi: 10.1016/j.cell.2020.07.007. [ This elegant study identifies distinct electrical oscillation patterns in neuroendocrine dopamine neurons as the basis of species-specific paternal strategies in male mice and rats ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R19] 19.Jamieson BB, et al. Overnight circuit remodelling drives juvenile alloparental care. 2025 doi: 10.1101/2025.06.13.659524. Preprint at. [DOI] [Google Scholar]

[R20] 20.Rogers FD, Bales KL. Mothers, Fathers, and Others: Neural Substrates of Parental Care. Trends Neurosci. 2019;42:552–562. doi: 10.1016/j.tins.2019.05.008. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R21] 21.Rilling JK, Mascaro JS. The neurobiology of fatherhood. Curr Opin Psychol. 2017;15:26–32. doi: 10.1016/j.copsyc.2017.02.013. [DOI] [PubMed] [Google Scholar]

[R22] 22.Clutton-Brock T. Cooperation between non-kin in animal societies. Nature. 2009;462:51–57. doi: 10.1038/nature08366. [DOI] [PubMed] [Google Scholar]

[R23] 23.Lukas D, Clutton-Brock TH. The Evolution of Social Monogamy in Mammals. Science. 2013;341:526–530. doi: 10.1126/science.1238677. [DOI] [PubMed] [Google Scholar]

[R24] 24.Hrdy SB. Mothers and Others: The Evolutionary Origins of Mutual Understanding. Vol. 422 Harvard University Press; Cambridge, MA, US: 2009. [Google Scholar]

[R25] 25.Kleiman DG. Monogamy in Mammals. Q Rev Biol. 1977;52:39–69. doi: 10.1086/409721. [DOI] [PubMed] [Google Scholar]

[R26] 26.Saltzman W, et al. Paternal Care in Biparental Rodents: Intra- and Inter-individual Variation. Integr Comp Biol. 2017;57:589–602. doi: 10.1093/icb/icx047. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R27] 27.Lukas D, Clutton-Brock T. Climate and the distribution of cooperative breeding in mammals. R Soc Open Sci. 2017;4:160897. doi: 10.1098/rsos.160897. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R28] 28.Hrdy SB, Burkart JM. The emergence of emotionally modern humans: implications for language and learning. Philos Trans R Soc B Biol Sci. 2020;375:20190499. doi: 10.1098/rstb.2019.0499. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R29] 29.Beach FA, Jaynes J. Studies of maternal retrieving in rats. III. Sensory cues involved in the lactating female’s response to her young. Behaviour. 1956;10:104–125. [Google Scholar]

[R30] 30.Rymer TL. The Role of Olfactory Genes in the Expression of Rodent Paternal Care Behavior. Genes. 2020;11:292. doi: 10.3390/genes11030292. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R31] 31.Kohl J, Autry AE, Dulac C. The neurobiology of parenting: A neural circuit perspective. BioEssays. 2017;39:1–11. doi: 10.1002/bies.201600159. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R32] 32.Isogai Y, et al. Multisensory Logic of Infant-Directed Aggression by Males. Cell. 2018;175:1827–1841.:e17. doi: 10.1016/j.cell.2018.11.032. [ This study finds that combined detection of chemosensory and tactile cues elicits pup-directed aggression in male mice ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R33] 33.Noirot E. Selective priming of maternal responses by auditory and olfactory cues from mouse pups. Dev Psychobiol. 1969;2:273–276. doi: 10.1002/dev.420020413. [DOI] [PubMed] [Google Scholar]

[R34] 34.Marlin BJ, Mitre M, D’amour JA, Chao MV, Froemke RC. Oxytocin enables maternal behaviour by balancing cortical inhibition. Nature. 2015;520:499–504. doi: 10.1038/nature14402. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R35] 35.Schiavo JK, et al. Innate and plastic mechanisms for maternal behaviour in auditory cortex. Nature. 2020;587:426–431. doi: 10.1038/s41586-020-2807-6. [ This study finds that experience-dependent maternal behaviour in mice results from interactions between intrinsic mechanisms and experience-dependent plasticity in the auditory cortex ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R36] 36.Tasaka G, et al. Genetic tagging of active neurons in auditory cortex reveals maternal plasticity of coding ultrasonic vocalizations. Nat Commun. 2018;9:871. doi: 10.1038/s41467-018-03183-2. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R37] 37.Tasaka G, et al. The Temporal Association Cortex Plays a Key Role in Auditory-Driven Maternal Plasticity. Neuron. 2020;107:566–579.:e7. doi: 10.1016/j.neuron.2020.05.004. [ This study identifies the temporal association cortex as a key cortical hub for processing pup vocalisations and shows that maternal experience enhances the discriminability of pup calls in this area ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R38] 38.Valtcheva S, et al. Neural circuitry for maternal oxytocin release induced by infant cries. Nature. 2023;621:788–795. doi: 10.1038/s41586-023-06540-4. [ Using in vivo recordings, this elegant study discovers that oxytocin neurons respond to pup vocalisations and that their activation is gated by disinhibition from the posterior intralaminar thalamus ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R39] 39.LeMessurier AM, Agha AA, Stephens J, Froemke RC. Auditory corticofugal circuits drive vocalization-guided maternal behavior. BioRxiv Prepr Serv Biol. 2023:2023.09.13.557594. doi: 10.1101/2023.09.13.557594. [DOI] [Google Scholar]

[R40] 40.Matsuo T, et al. Genetic dissection of pheromone processing reveals main olfactory system-mediated social behaviors in mice. Proc Natl Acad Sci U S A. 2015;112:E311–320. doi: 10.1073/pnas.1416723112. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R41] 41.Wang Z, Storm DR. Maternal Behavior is Impaired in Female Mice Lacking Type 3 Adenylyl Cyclase. Neuropsychopharmacology. 2011;36:772–781. doi: 10.1038/npp.2010.211. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R42] 42.Zarrow M, Gandelman R, Denenberg VH. Lack of nest building and maternal behavior in the mouse following olfactory bulb removal. Horm Behav. 1971;2:227–238. [Google Scholar]

[R43] 43.Gandelman R, Zarrow MX, Denenberg VH, Myers M. Olfactory Bulb Removal Eliminates Maternal Behavior in the Mouse. Science. 1971;171:210–211. doi: 10.1126/science.171.3967.210. [DOI] [PubMed] [Google Scholar]

[R44] 44.Canteras NS, Simerly RB, Swanson LW. Organization of projections from the medial nucleus of the amygdala: a PHAL study in the rat. J Comp Neurol. 1995;360:213–245. doi: 10.1002/cne.903600203. [DOI] [PubMed] [Google Scholar]

[R45] 45.Raam T, Hong W. Organization of neural circuits underlying social behavior: A consideration of the medial amygdala. Curr Opin Neurobiol. 2021;68:124–136. doi: 10.1016/j.conb.2021.02.008. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R46] 46.Chen PB, et al. Sexually Dimorphic Control of Parenting Behavior by the Medial Amygdala. Cell. 2019;176:1206–1221.:e18. doi: 10.1016/j.cell.2019.01.024. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R47] 47.Kohl J, et al. Functional circuit architecture underlying parental behaviour. Nature. 2018;556:326–331. doi: 10.1038/s41586-018-0027-0. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R48] 48.Kohl J, Dulac C. Neural control of parental behaviors. Curr Opin Neurobiol. 2018;49:116–122. doi: 10.1016/j.conb.2018.02.002. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R49] 49.Keller M, Perrin G, Meurisse M, Ferreira G, Lévy F. Cortical and medial amygdala are both involved in the formation of olfactory offspring memory in sheep. Eur J Neurosci. 2004;20:3433–3441. doi: 10.1111/j.1460-9568.2004.03812.x. [DOI] [PubMed] [Google Scholar]

[R50] 50.Nowak R, Keller M, Lévy F. Mother–Young Relationships in Sheep: A Model for a Multidisciplinary Approach of the Study of Attachment in Mammals. J Neuroendocrinol. 2011;23:1042–1053. doi: 10.1111/j.1365-2826.2011.02205.x. [DOI] [PubMed] [Google Scholar]

[R51] 51.Nowlan AC, et al. Multisensory integration of social signals by a pathway from the basal amygdala to the auditory cortex in maternal mice. Curr Biol. 2025;35:36–49.:e4. doi: 10.1016/j.cub.2024.10.078. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R52] 52.Tachikawa KS, Yoshihara Y, Kuroda KO. Behavioral Transition from Attack to Parenting in Male Mice: A Crucial Role of the Vomeronasal System. J Neurosci. 2013;33:5120–5126. doi: 10.1523/JNEUROSCI.2364-12.2013. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R53] 53.Wu Z, Autry AE, Bergan JF, Watabe-Uchida M, Dulac CG. Galanin neurons in the medial preoptic area govern parental behaviour. Nature. 2014;509:325–330. doi: 10.1038/nature13307. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R54] 54.Prakash N, Abu Irqeba A, Corbin JG. Development and function of the medial amygdala. Trends Neurosci. 2025;48:22–32. doi: 10.1016/j.tins.2024.11.004. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R55] 55.Keshavarzi S, Sullivan RKP, Ianno DJ, Sah P. Functional properties and projections of neurons in the medial amygdala. J Neurosci Off J Soc Neurosci. 2014;34:8699–8715. doi: 10.1523/JNEUROSCI.1176-14.2014. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R56] 56.Sheehan T, Paul M, Amaral E, Numan MJ, Numan M. Evidence that the medial amygdala projects to the anterior/ventromedial hypothalamic nuclei to inhibit maternal behavior in rats. Neuroscience. 2001;106:341–356. doi: 10.1016/s0306-4522(01)00286-x. [DOI] [PubMed] [Google Scholar]

[R57] 57.Xie Z, et al. Mechanically evoked defensive attack is controlled by GABAergic neurons in the anterior hypothalamic nucleus. Nat Neurosci. 2022;25:72–85. doi: 10.1038/s41593-021-00985-4. [DOI] [PubMed] [Google Scholar]

[R58] 58.Wang L, Chen IZ, Lin D. Collateral Pathways from the Ventromedial Hypothalamus Mediate Defensive Behaviors. Neuron. 2015;85:1344–1358. doi: 10.1016/j.neuron.2014.12.025. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R59] 59.Lischinsky JE, et al. Transcriptionally defined amygdala subpopulations play distinct roles in innate social behaviors. Nat Neurosci. 2023;26:2131–2146. doi: 10.1038/s41593-023-01475-5. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R60] 60.Kenyon P, Cronin P, Keeble S. Disruption of maternal retrieving by perioral anesthesia. Physiol Behav. 1981;27:313–321. doi: 10.1016/0031-9384(81)90274-2. [DOI] [PubMed] [Google Scholar]

[R61] 61.Kenyon P, Cronin P, Keeble S. Role of the infraorbital nerve in retrieving behavior in lactating rats. Behav Neurosci. 1983;97:255–269. doi: 10.1037//0735-7044.97.2.255. [DOI] [PubMed] [Google Scholar]

[R62] 62.Salay LD, Tsao DY, Anderson DJ. A circuit that integrates drive state and social contact to gate mating. Nature. 2025:1–10. doi: 10.1038/s41586-025-09327-x. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R63] 63.Cservenák M, et al. Tuberoinfundibular Peptide of 39 Residues Is Activated during Lactation and Participates in the Suckling-Induced Prolactin Release in Rat. Endocrinology. 2010;151:5830–5840. doi: 10.1210/en.2010-0767. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R64] 64.Cservenák M, et al. Thalamic neuropeptide mediating the effects of nursing on lactation and maternal motivation. Psychoneuroendocrinology. 2013;38:3070–3084. doi: 10.1016/j.psyneuen.2013.09.004. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R65] 65.Cservenák M, et al. A Thalamo-Hypothalamic Pathway That Activates Oxytocin Neurons in Social Contexts in Female Rats. Endocrinology. 2017;158:335–348. doi: 10.1210/en.2016-1645. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R66] 66.Scott N, Prigge M, Yizhar O, Kimchi T. A sexually dimorphic hypothalamic circuit controls maternal care and oxytocin secretion. Nature. 2015;525:519–522. doi: 10.1038/nature15378. [DOI] [PubMed] [Google Scholar]

[R67] 67.Cservenák M, et al. Maternally involved galanin neurons in the preoptic area of the rat. Brain Struct Funct. 2017;222:781–798. doi: 10.1007/s00429-016-1246-5. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R68] 68.Rosselet C, Zennou-Azogui Y, Xerri C. Nursing-Induced Somatosensory Cortex Plasticity: Temporally Decoupled Changes in Neuronal Receptive Field Properties Are Accompanied by Modifications in Activity-Dependent Protein Expression. J Neurosci. 2006;26:10667–10676. doi: 10.1523/JNEUROSCI.3253-06.2006. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R69] 69.Okabe S, et al. Pup odor and ultrasonic vocalizations synergistically stimulate maternal attention in mice. Behav Neurosci. 2013;127:432–438. doi: 10.1037/a0032395. [DOI] [PubMed] [Google Scholar]

[R70] 70.Lecca S, et al. A neural substrate for negative affect dictates female parental behavior. Neuron. 2023;111:1094–1103.:e8. doi: 10.1016/j.neuron.2023.01.003. [ This study demonstrates that pup vocalisations carry negative valence in virgin female mice ] [DOI] [PubMed] [Google Scholar]

[R71] 71.Carta I, et al. Sex-specific hypothalamic neural projection activity drives caregiving in mice. Nat Commun. 2025;16:4116. doi: 10.1038/s41467-025-59352-7. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R72] 72.Gómora-Arrati P, Dominguez G, Ågmo A. GABA Receptors in the Medial Preoptic Area Modulate the Onset of Oestradiol-Induced Maternal Behaviour in Hysterectomised-Ovariectomised, Pregnant Rats. J Neuroendocrinol. 2016;28:jne.12434. doi: 10.1111/jne.12434. [DOI] [PubMed] [Google Scholar]

[R73] 73.Numan M, Numan M. A lesion and neuroanatomical tract-tracing analysis of the role of the bed nucleus of the stria terminalis in retrieval behavior and other aspects of maternal responsiveness in rats. Dev Psychobiol. 1996;29:23–51. doi: 10.1002/(SICI)1098-2302(199601)29:1<23::AID-DEV2>3.0.CO;2-O. [DOI] [PubMed] [Google Scholar]

[R74] 74.Perrin G, Meurisse M, Levy F. Inactivation of the medial preoptic area or the bed nucleus of the stria terminalis differentially disrupts maternal behavior in sheep. Horm Behav. 2007;52:461–473. doi: 10.1016/j.yhbeh.2007.06.010. [DOI] [PubMed] [Google Scholar]

[R75] 75.Benedict J, Cudmore RH, Oden D, Spruell A, Linden DJ. The Lateral Habenula Is Necessary for Maternal Behavior in the Naturally Parturient Primiparous Mouse Dam. eneuro. 2025;12:ENEURO.0092-24.2024. doi: 10.1523/ENEURO.0092-24.2024. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R76] 76.Numan M. Medial preoptic area and maternal behavior in the female rat. J Comp Physiol Psychol. 1974;87:746–759. doi: 10.1037/h0036974. [DOI] [PubMed] [Google Scholar]

[R77] 77.Jacobson CD, Terkel J, Gorski RA, Sawyer CH. Effects of small medial preoptic area lesions on maternal behavior: Retreiving and nest building in the rat. Brain Res. 1980;194:471–478. doi: 10.1016/0006-8993(80)91226-3. [DOI] [PubMed] [Google Scholar]

[R78] 78.Tsuneoka Y, et al. Functional, anatomical, and neurochemical differentiation of medial preoptic area subregions in relation to maternal behavior in the mouse. J Comp Neurol. 2013;521:1633–1663. doi: 10.1002/cne.23251. [DOI] [PubMed] [Google Scholar]

[R79] 79.Tsuneoka Y, et al. Distinct preoptic-BST nuclei dissociate paternal and infanticidal behavior in mice. EMBO J. 2015;34:2652–2670. doi: 10.15252/embj.201591942. [ This study finds that central MPOA lesions promote infanticidal behavior in fathers, while lesions of the rhomboid nucleus of the BNST inhibit infanticide in virgin males ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R80] 80.Kurachi T, et al. Behavioral competition between infant care and sexual behavior in male but not female common marmosets. Neurosci Res. 2025;216:104900. doi: 10.1016/j.neures.2025.04.003. [DOI] [PubMed] [Google Scholar]

[R81] 81.Bridges RS. Neuroendocrine regulation of maternal behavior. Front Neuroendocrinol. 2015;36:178–196. doi: 10.1016/j.yfrne.2014.11.007. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R82] 82.Fang Y-Y, Yamaguchi T, Song SC, Tritsch NX, Lin D. A Hypothalamic Midbrain Pathway Essential for Driving Maternal Behaviors. Neuron. 2018;98:192–207.:e10. doi: 10.1016/j.neuron.2018.02.019. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R83] 83.Xie Y, Huang L, Corona A, Pagliaro AH, Shea SD. A dopaminergic reward prediction error signal shapes maternal behavior in mice. Neuron. 2023;111:557–570.:e7. doi: 10.1016/j.neuron.2022.11.019. [ This elegant study establishes that dopaminergic VTA neurons refine stereotyped pup retrieval behaviour by a reinforcement learning mechanism ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R84] 84.Moffitt JR, et al. Molecular, spatial, and functional single-cell profiling of the hypothalamic preoptic region. Science. 2018;362:eaau5324. doi: 10.1126/science.aau5324. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R85] 85.Wei Y-C, et al. Medial preoptic area in mice is capable of mediating sexually dimorphic behaviors regardless of gender. Nat Commun. 2018;9:279. doi: 10.1038/s41467-017-02648-0. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R86] 86.Zhang Z, et al. Estrogen-sensitive medial preoptic area neurons coordinate torpor in mice. Nat Commun. 2020;11:6378. doi: 10.1038/s41467-020-20050-1. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R87] 87.Yoshihara C, et al. Calcitonin receptor signaling in the medial preoptic area enables risk-taking maternal care. Cell Rep. 2021;35:109204. doi: 10.1016/j.celrep.2021.109204. [DOI] [PubMed] [Google Scholar]

[R88] 88.Nair A, et al. The neural computation of affective internal states in the hypothalamus: A dynamical systems perspective. Neuron. 2025;113:3887–3907. doi: 10.1016/j.neuron.2025.11.003. [DOI] [PubMed] [Google Scholar]

[R89] 89.Autry AE, et al. Urocortin-3 neurons in the mouse perifornical area promote infant-directed neglect and aggression. eLife. 2021;10:e64680. doi: 10.7554/eLife.64680. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R90] 90.Sato K, et al. Amygdalohippocampal Area Neurons That Project to the Preoptic Area Mediate Infant-Directed Attack in Male Mice. J Neurosci. 2020;40:3981–3994. doi: 10.1523/JNEUROSCI.0438-19.2020. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R91] 91.Mei L, et al. The neural control of infanticide and parental behaviors in male mice. 2026 doi: 10.64898/2026.03.12.711470. Preprint at. [DOI] [Google Scholar]

[R92] 92.Lee A, Clancy S, Fleming AS. Mother rats bar-press for pups: effects of lesions of the mpoa and limbic sites on maternal behavior and operant responding for pup-reinforcement. Behav Brain Res. 1999;100:15–31. doi: 10.1016/s0166-4328(98)00109-0. [DOI] [PubMed] [Google Scholar]

[R93] 93.Mattson BJ, Morrell JI. Preference for cocaine-versus pup-associated cues differentially activates neurons expressing either Fos or cocaine- and amphetamine-regulated transcript in lactating, maternal rodents. Neuroscience. 2005;135:315–328. doi: 10.1016/j.neuroscience.2005.06.045. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R94] 94.Feldman R. The adaptive human parental brain: implications for children’s social development. Trends Neurosci. 2015;38:387–399. doi: 10.1016/j.tins.2015.04.004. [DOI] [PubMed] [Google Scholar]

[R95] 95.Day KR, Shea SD. Dopamine dynamics underlying maternal motivation and reward. Neurosci Res. 2025;218:104928. doi: 10.1016/j.neures.2025.104928. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R96] 96.Hansen S, Harthon C, Wallin E, Löfberg L, Svensson K. Mesotelencephalic dopamine system and reproductive behavior in the female rat: Effects of ventral tegmental 6-hydroxydopamine lesions on maternal and sexual responsiveness. Behav Neurosci. 1991;105:588–598. doi: 10.1037//0735-7044.105.4.588. [DOI] [PubMed] [Google Scholar]

[R97] 97.Hansen S. Maternal behavior of female rats with 6-OHDA lesions in the ventral striatum: Characterization of the pup retrieval deficit. Physiol Behav. 1994;55:615–620. doi: 10.1016/0031-9384(94)90034-5. [DOI] [PubMed] [Google Scholar]

[R98] 98.Numan M, Stolzenberg DS, Dellevigne AA, Correnti CM, Numan MJ. Temporary inactivation of ventral tegmental area neurons with either muscimol or baclofen reversibly disrupts maternal behavior in rats through different underlying mechanisms. Behav Neurosci. 2009;123:740–751. doi: 10.1037/a0016204. [DOI] [PubMed] [Google Scholar]

[R99] 99.Keer SE, Stern JM. Dopamine Receptor Blockade in the Nucleus Accumbens Inhibits Maternal Retrieval and Licking, but Enhances Nursing Behavior in Lactating Rats. Physiol Behav. 1999;67:659–669. doi: 10.1016/s0031-9384(99)00116-x. [DOI] [PubMed] [Google Scholar]

[R100] 100.Li M, Fleming AS. Differential involvement of nucleus accumbens shell and core subregions in maternal memory in postpartum female rats. Behav Neurosci. 2003;117:426–445. doi: 10.1037/0735-7044.117.3.426. [DOI] [PubMed] [Google Scholar]

[R101] 101.Champagne FA, et al. Variations in Nucleus Accumbens Dopamine Associated with Individual Differences in Maternal Behavior in the Rat. J Neurosci. 2004;24:4113–4123. doi: 10.1523/JNEUROSCI.5322-03.2004. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R102] 102.Alsina-Llanes M, Olazábal DE. NMDA- and 6-OHDA-induced Lesions in the Nucleus Accumbens Differently Affect Maternal and Infanticidal Behavior in Pup-naïve Female and Male Mice. Neuroscience. 2024;539:35–50. doi: 10.1016/j.neuroscience.2023.12.009. [DOI] [PubMed] [Google Scholar]

[R103] 103.Strathearn L, Li J, Fonagy P, Montague PR. What’s in a Smile? Maternal Brain Responses to Infant Facial Cues. Pediatrics. 2008;122:40–51. doi: 10.1542/peds.2007-1566. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R104] 104.Strathearn L, Fonagy P, Amico J, Montague PR. Adult Attachment Predicts Maternal Brain and Oxytocin Response to Infant Cues. Neuropsychopharmacology. 2009;34:2655–2666. doi: 10.1038/npp.2009.103. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R105] 105.Tasaka G-I, et al. Orbitofrontal cortex influences dopamine dynamics associated with alloparental behavioral acquisition in female mice. Sci Adv. 2025;11:eadr4620. doi: 10.1126/sciadv.adr4620. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R106] 106.Stempel AV. A conserved brainstem region for instinctive behaviour control: The vertebrate periaqueductal gray. Curr Opin Neurobiol. 2024;86:102878. doi: 10.1016/j.conb.2024.102878. [DOI] [PubMed] [Google Scholar]

[R107] 107.Bandler R, Shipley MT. Columnar organization in the midbrain periaqueductal gray: modules for emotional expression? Trends Neurosci. 1994;17:379–389. doi: 10.1016/0166-2236(94)90047-7. [DOI] [PubMed] [Google Scholar]

[R108] 108.Lonstein JS, Stern JM. Role of the Midbrain Periaqueductal Gray in Maternal Nurturance and Aggression: c-fos and Electrolytic Lesion Studies in Lactating Rats. J Neurosci. 1997;17:3364–3378. doi: 10.1523/JNEUROSCI.17-09-03364.1997. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R109] 109.Lonstein JS, Stern JM. Site and behavioral specificity of periaqueductal gray lesions on postpartum sexual, maternal, and aggressive behaviors in rats. Brain Res. 1998;804:21–35. doi: 10.1016/s0006-8993(98)00642-8. [DOI] [PubMed] [Google Scholar]

[R110] 110.Lonstein JS, Simmons DA, Stern JM. Functions of the caudal periaqueductal gray in lactating rats: Kyphosis, lordosis, maternal aggression, and fearfulness. Behav Neurosci. 1998;112:1502–1518. doi: 10.1037//0735-7044.112.6.1502. [DOI] [PubMed] [Google Scholar]

[R111] 111.Zhang G-W, et al. Medial preoptic area antagonistically mediates stress-induced anxiety and parental behavior. Nat Neurosci. 2021;24:516–528. doi: 10.1038/s41593-020-00784-3. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R112] 112.Zhu Z, et al. A substantia innominata-midbrain circuit controls a general aggressive response. Neuron. 2021;109:1540–1553.:e9. doi: 10.1016/j.neuron.2021.03.002. [DOI] [PubMed] [Google Scholar]

[R113] 113.Numan M, Numan MJ. Preoptic-brainstem connections and maternal behavior in rats. Behav Neurosci. 1991;105:1013–1029. doi: 10.1037//0735-7044.105.6.1013. [DOI] [PubMed] [Google Scholar]

[R114] 114.Cohen L, Rothschild G, Mizrahi A. Multisensory Integration of Natural Odors and Sounds in the Auditory Cortex. Neuron. 2011;72:357–369. doi: 10.1016/j.neuron.2011.08.019. [DOI] [PubMed] [Google Scholar]

[R115] 115.Afonso VM, Sison M, Lovic V, Fleming AS. Medial prefrontal cortex lesions in the female rat affect sexual and maternal behavior and their sequential organization. Behav Neurosci. 2007;121:515–526. doi: 10.1037/0735-7044.121.3.515. [DOI] [PubMed] [Google Scholar]

[R116] 116.Febo M, Felix-Ortiz AC, Johnson TR. Inactivation or inhibition of neuronal activity in the medial prefrontal cortex largely reduces pup retrieval and grouping in maternal rats. Brain Res. 2010;1325:77–88. doi: 10.1016/j.brainres.2010.02.027. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R117] 117.Wu K, et al. Experience-dependent maternal defense behavior mediated by profrontal cortical projections to the medial preoptic area in mice. Nat Commun. 2025;16:7664. doi: 10.1038/s41467-025-63062-5. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R118] 118.Swain JE, et al. Approaching the biology of human parental attachment: Brain imaging, oxytocin and coordinated assessments of mothers and fathers. Brain Res. 2014;1580:78–101. doi: 10.1016/j.brainres.2014.03.007. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R119] 119.Musser ED, Kaiser-Laurent H, Ablow JC. The neural correlates of maternal sensitivity: An fMRI study. Dev Cogn Neurosci. 2012;2:428–436. doi: 10.1016/j.dcn.2012.04.003. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R120] 120.Gandelman R. The ontogeny of maternal responsiveness in female Rockland-Swiss albino mice. Horm Behav. 1973;4:257–268. doi: 10.1016/0018-506x(73)90010-x. [DOI] [PubMed] [Google Scholar]

[R121] 121.Gandelman R. The development of cannibalism in male Rockland-Swiss mice and the influence of olfactory bulb removal. Dev Psychobiol. 1973;6:159–164. doi: 10.1002/dev.420060210. [DOI] [PubMed] [Google Scholar]

[R122] 122.Rosenblatt JS. Nonhormonal Basis of Maternal Behavior in the Rat. Science. 1967;156:1512–1514. doi: 10.1126/science.156.3781.1512. [DOI] [PubMed] [Google Scholar]

[R123] 123.Husain BFA, Kohl J. Plasticity of the parental brain. Curr Opin Neurobiol. 2025;95:103129. doi: 10.1016/j.conb.2025.103129. [DOI] [PubMed] [Google Scholar]

[R124] 124.Servin-Barthet C, et al. The transition to motherhood: linking hormones, brain and behaviour. Nat Rev Neurosci. 2023;24:605–619. doi: 10.1038/s41583-023-00733-6. [DOI] [PubMed] [Google Scholar]

[R125] 125.Hoekzema E, et al. Pregnancy leads to long-lasting changes in human brain structure. Nat Neurosci. 2017;20:287–296. doi: 10.1038/nn.4458. [DOI] [PubMed] [Google Scholar]

[R126] 126.Kenkel WM, Perkeybile AM, Carter CS. The neurobiological causes and effects of alloparenting. Dev Neurobiol. 2017;77:214–232. doi: 10.1002/dneu.22465. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R127] 127.Feldman R, Braun K, Champagne FA. The neural mechanisms and consequences of paternal caregiving. Nat Rev Neurosci. 2019;20:205–224. doi: 10.1038/s41583-019-0124-6. [DOI] [PubMed] [Google Scholar]

[R128] 128.Bridges RS, Zarrow MX, Goldman BD, Denenberg VH. A developmental study of maternal responsiveness in the rat. Physiol Behav. 1974;12:149–151. doi: 10.1016/0031-9384(74)90082-1. [DOI] [PubMed] [Google Scholar]

[R129] 129.Mayer AD, Rosenblatt JS. Ontogeny of maternal behavior in the laboratory rat: Early origins in 18-to 27-day-old young. Dev Psychobiol. 1979;12:407–424. doi: 10.1002/dev.420120502. [DOI] [PubMed] [Google Scholar]

[R130] 130.Roberts RL, Miller AK, Taymans SE, Carter CS. Role of social and endocrine factors in alloparental behavior of prairie voles (Microtus ochrogaster) Can J Zool. 1998;76:1862–1868. [Google Scholar]

[R131] 131.Olazábal DE, Young LJ. Variability in “spontaneous” maternal behavior is associated with anxiety-like behavior and affiliation in Naïve juvenile and adult female prairie voles (Microtus ochrogaster) Dev Psychobiol. 2005;47:166–178. doi: 10.1002/dev.20077. [DOI] [PubMed] [Google Scholar]

[R132] 132.Kaplan HS, et al. Sensory input, sex and function shape hypothalamic cell type development. Nature. 2025;647:157–168. doi: 10.1038/s41586-025-08603-0. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R133] 133.Kercmar J, Snoj T, Tobet SA, Majdic G. Gonadectomy prior to puberty decreases normal parental behavior in adult mice. Horm Behav. 2014;66:667–673. doi: 10.1016/j.yhbeh.2014.09.007. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R134] 134.Fukui K, Uki H, Minami M, Amano T. Effect of gonadal steroid hormone levels during pubertal development on social behavior of adult mice toward pups and synaptic transmission in the rhomboid nucleus of the bed nucleus of the stria terminalis. Neurosci Lett. 2019;708:134357. doi: 10.1016/j.neulet.2019.134357. [DOI] [PubMed] [Google Scholar]

[R135] 135.Weissner B, Sheard N. Maternal behaviour in the rat. Endocrinology. 1933;17:590. [Google Scholar]

[R136] 136.Leblond CP. Extra-Hormonal Factors in Maternal Behavior. Exp Biol Med. 1938;38:66–70. [Google Scholar]

[R137] 137.Alsina-Llanes M, Lenzi B, Olazábal DE. Repeated non-contact exposure to pups inhibits infanticidal and facilitates paternal behavior in virgin adult male mice (C57BL6) Physiol Behav. 2025;295:114885. doi: 10.1016/j.physbeh.2025.114885. [DOI] [PubMed] [Google Scholar]

[R138] 138.Cai W, et al. Involvement of the dopamine system in paternal behavior induced by repeated pup exposure in virgin male ICR mice. Behav Brain Res. 2021;415:113519. doi: 10.1016/j.bbr.2021.113519. [DOI] [PubMed] [Google Scholar]

[R139] 139.Stolzenberg DS, Rissman EF. Oestrogen-Independent, Experience-Induced Maternal Behaviour in Female Mice: Oestradiol-independent maternal behaviour. J Neuroendocrinol. 2011;23:345–354. doi: 10.1111/j.1365-2826.2011.02112.x. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R140] 140.Noirot E. Changes in responsiveness to young in the adult mouse. I. The problematical effect of hormones. Anim Behav. 1964;12:52–58. [Google Scholar]

[R141] 141.Noirot E. Changes in responsiveness to young in the adult mouse. IV. The effect of an initial contact with a strong stimulus. Anim Behav. 1964;12:442–445. [Google Scholar]

[R142] 142.Cohen J, Bridges RS. Retention of maternal behavior in nulliparous and primiparous rats: Effects of duration of previous maternal experience. J Comp Physiol Psychol. 1981;95:450–459. [Google Scholar]

[R143] 143.Bridges RS, Scanlan VF. Maternal memory in adult, nulliparous rats: Effects of testing interval on the retention of maternal behavior. Dev Psychobiol. 2005;46:13–18. doi: 10.1002/dev.20038. [DOI] [PubMed] [Google Scholar]

[R144] 144.Okabe S, et al. Pup exposure facilitates retrieving behavior via the oxytocin neural system in female mice. Psychoneuroendocrinology. 2017;79:20–30. doi: 10.1016/j.psyneuen.2017.01.036. [DOI] [PubMed] [Google Scholar]

[R145] 145.Bair-Marshall C, Cassel N, Agha A, Bkhiet M, Froemke R. Neural circuit plasticity transforms infant neglect into maternal care. 2025 doi: 10.21203/rs.3.rs-5983736/v1. Preprint at. [DOI] [Google Scholar]

[R146] 146.Swart JM, Grattan DR, Ladyman SR, Brown RSE. Pups and prolactin are rewarding to virgin female and pregnant mice. J Neuroendocrinol. 2023;35:e13232. doi: 10.1111/jne.13232. [DOI] [PubMed] [Google Scholar]

[R147] 147.Salais-López H, et al. Maternal Motivation: Exploring the Roles of Prolactin and Pup Stimuli. Neuroendocrinology. 2021;111:805–830. doi: 10.1159/000510038. [DOI] [PubMed] [Google Scholar]

[R148] 148.Quinones-Laracuente K, et al. A collicular-hypothalamic pathway for social visual awareness. BioRxiv Prepr Serv Biol. 2026:2026.04.17.719268. doi: 10.64898/2026.04.17.719268. [DOI] [Google Scholar]

[R149] 149.Rawn SM, et al. Pregnancy Hyperglycemia in Prolactin Receptor Mutant, but Not Prolactin Mutant, Mice and Feeding-Responsive Regulation of Placental Lactogen Genes Implies Placental Control of Maternal Glucose Homeostasis. Biol Reprod. 2015;93(75):1–12. doi: 10.1095/biolreprod.115.132431. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R150] 150.Soares MJ. The prolactin and growth hormone families: Pregnancy-specific hormones/cytokines at the maternal-fetal interface. Reprod Biol Endocrinol. 2004;2:51. doi: 10.1186/1477-7827-2-51. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R151] 151.Brown RSE, et al. Prolactin action in the medial preoptic area is necessary for postpartum maternal nursing behavior. Proc Natl Acad Sci. 2017;114:10779–10784. doi: 10.1073/pnas.1708025114. [ This study establishes a critical role for prolactin / placental lactogen action in the MPOA for maternal caregiving behaviour in mice ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R152] 152.Georgescu T, Swart JM, Grattan DR, Brown RSE. The Prolactin Family of Hormones as Regulators of Maternal Mood and Behavior. Front Glob Womens Health. 2021;2:767467. doi: 10.3389/fgwh.2021.767467. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R153] 153.Smiley KO, Brown RSE, Grattan DR. Prolactin Action Is Necessary for Parental Behavior in Male Mice. J Neurosci. 2022;42:8308–8327. doi: 10.1523/JNEUROSCI.0558-22.2022. [ This study finds that prolactin action in the forebrain is critical for the establishment of paternal behaviour in male mice ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R154] 154.Francis K, Meddle SL, Bishop VR, Russell JA. Progesterone receptor expression in the pregnant and parturient rat hypothalamus and brainstem. Brain Res. 2002;927:18–26. doi: 10.1016/s0006-8993(01)03318-2. [DOI] [PubMed] [Google Scholar]

[R155] 155.Doerr HK, Siegel HI, Rosenblatt JS. Effects of progesterone withdrawal and estrogen on maternal behavior in nulliparous rats. Behav Neural Biol. 1981;32:35–44. doi: 10.1016/s0163-1047(81)90242-9. [DOI] [PubMed] [Google Scholar]

[R156] 156.Murakami G. Distinct Effects of Estrogen on Mouse Maternal Behavior: The Contribution of Estrogen Synthesis in the Brain. PLOS ONE. 2016;11:e0150728. doi: 10.1371/journal.pone.0150728. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R157] 157.Rosenblatt JS, Wagner CK, Morrell JI. Hormonal priming and triggering of maternal behavior in the rat with special reference to the relations between estrogen receptor binding and ER mRNA in specific brain regions. Psychoneuroendocrinology. 1994;19:543–552. doi: 10.1016/0306-4530(94)90039-6. [DOI] [PubMed] [Google Scholar]

[R158] 158.Glynn LM, Davis EP, Sandman CA, Goldberg WA. Gestational hormone profiles predict human maternal behavior at 1-year postpartum. Horm Behav. 2016;85:19–25. doi: 10.1016/j.yhbeh.2016.07.002. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R159] 159.Ribeiro AC, et al. siRNA silencing of estrogen receptor-α expression specifically in medial preoptic area neurons abolishes maternal care in female mice. Proc Natl Acad Sci. 2012;109:16324–16329. doi: 10.1073/pnas.1214094109. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R160] 160.Ogawa S, et al. Roles of Estrogen Receptor-α Gene Expression in Reproduction-Related Behaviors in Female Mice. Endocrinology. 1998;139:5070–5081. doi: 10.1210/endo.139.12.6357. [DOI] [PubMed] [Google Scholar]

[R161] 161.Catanese MC, Vandenberg LN. Bisphenol S (BPS) alters maternal behavior and brain in mice exposed during pregnancy/lactation and their daughters. Endocrinology. 2016:en.2016-1723. doi: 10.1210/en.2016-1723. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R162] 162.Champagne FA, et al. Maternal Care Associated with Methylation of the Estrogen Receptor-α1b Promoter and Estrogen Receptor-α Expression in the Medial Preoptic Area of Female Offspring. Endocrinology. 2006;147:2909–2915. doi: 10.1210/en.2005-1119. [DOI] [PubMed] [Google Scholar]

[R163] 163.Champagne FA, Weaver ICG, Diorio J, Sharma S, Meaney MJ. Natural Variations in Maternal Care Are Associated with Estrogen Receptor α Expression and Estrogen Sensitivity in the Medial Preoptic Area. Endocrinology. 2003;144:4720–4724. doi: 10.1210/en.2003-0564. [DOI] [PubMed] [Google Scholar]

[R164] 164.Topilko T, et al. Edinger-Westphal peptidergic neurons enable maternal preparatory nesting. Neuron. 2022;110:1385–1399.:e8. doi: 10.1016/j.neuron.2022.01.012. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R165] 165.Brunton PJ, Russell JA. The expectant brain: adapting for motherhood. Nat Rev Neurosci. 2008;9:11–25. doi: 10.1038/nrn2280. [DOI] [PubMed] [Google Scholar]

[R166] 166.Neumann I, Douglas AJ, Pittman QJ, Russell JA, Landgraf R. Oxytocin Released within the Supraoptic Nucleus of the Rat Brain by Positive Feedback Action is Involved in Parturition-Related Events. J Neuroendocrinol. 1996;8:227–233. doi: 10.1046/j.1365-2826.1996.04557.x. [DOI] [PubMed] [Google Scholar]

[R167] 167.Jurek B, Neumann ID. The Oxytocin Receptor: From Intracellular Signaling to Behavior. Physiol Rev. 2018;98:1805–1908. doi: 10.1152/physrev.00031.2017. [DOI] [PubMed] [Google Scholar]

[R168] 168.Menon R, Neumann ID. Detection, processing and reinforcement of social cues: regulation by the oxytocin system. Nat Rev Neurosci. 2023;24:761–777. doi: 10.1038/s41583-023-00759-w. [DOI] [PubMed] [Google Scholar]

[R169] 169.Meddle SL, Bishop VR, Gkoumassi E, Van Leeuwen FW, Douglas AJ. Dynamic Changes in Oxytocin Receptor Expression and Activation at Parturition in the Rat Brain. Endocrinology. 2007;148:5095–5104. doi: 10.1210/en.2007-0615. [DOI] [PubMed] [Google Scholar]

[R170] 170.Pedersen CA, Prange AJ. Induction of maternal behavior in virgin rats after intracerebroventricular administration of oxytocin. Proc Natl Acad Sci. 1979;76:6661–6665. doi: 10.1073/pnas.76.12.6661. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R171] 171.Pedersen CA, Caldwell JD, Johnson MF, Fort SA, Prange AJ. Oxytocin antiserum delays onset of ovarian steroid-induced maternal behavior. Neuropeptides. 1985;6:175–182. doi: 10.1016/0143-4179(85)90108-8. [DOI] [PubMed] [Google Scholar]

[R172] 172.Kendrick KM, Keverne EB, Baldwin BA. Intracerebroventricular Oxytocin Stimulates Maternal Behaviour in the Sheep. Neuroendocrinology. 1987;46:56–61. doi: 10.1159/000124796. [DOI] [PubMed] [Google Scholar]

[R173] 173.Fahrbach SE, Morrell JI, Pfaff DW. Possible Role for Endogenous Oxytocin in Estrogen-Facilitated Maternal Behavior in Rats. Neuroendocrinology. 1985;40:526–532. doi: 10.1159/000124125. [DOI] [PubMed] [Google Scholar]

[R174] 174.Boccia ML, Goursaud A-PS, Bachevalier J, Anderson KD, Pedersen CA. Peripherally administered non-peptide oxytocin antagonist, L368,899, accumulates in limbic brain areas: A new pharmacological tool for the study of social motivation in non-human primates. Horm Behav. 2007;52:344–351. doi: 10.1016/j.yhbeh.2007.05.009. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R175] 175.Nishimori K, et al. Oxytocin is required for nursing but is not essential for parturition or reproductive behavior. Proc Natl Acad Sci. 1996;93:11699–11704. doi: 10.1073/pnas.93.21.11699. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R176] 176.Takayanagi Y, et al. Pervasive social deficits, but normal parturition, in oxytocin receptor-deficient mice. Proc Natl Acad Sci. 2005;102:16096–16101. doi: 10.1073/pnas.0505312102. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R177] 177.Pedersen CA, Vadlamudi SV, Boccia ML, Amico JA. Maternal behavior deficits in nulliparous oxytocin knockout mice. Genes Brain Behav. 2006;5:274–281. doi: 10.1111/j.1601-183X.2005.00162.x. [DOI] [PubMed] [Google Scholar]

[R178] 178.Rich ME, deCárdenas EJ, Lee H-J, Caldwell HK. Impairments in the Initiation of Maternal Behavior in Oxytocin Receptor Knockout Mice. PLoS ONE. 2014;9:e98839. doi: 10.1371/journal.pone.0098839. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R179] 179.Tsuneoka Y, et al. Oxytocin Facilitates Allomaternal Behavior under Stress in Laboratory Mice. eneuro. 2022;9:ENEURO.0405-21.2022. doi: 10.1523/ENEURO.0405-21.2022. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R180] 180.Labriola J. Effects of Caesarean Delivery upon Maternal Behavior in Rats. Exp Biol Med. 1953;83:556–557. doi: 10.3181/00379727-83-20416. [DOI] [PubMed] [Google Scholar]

[R181] 181.Moltz H, Robbins D, Parks M. Caesarean delivery and maternal behavior of primiparous and multiparous rats. J Comp Physiol Psychol. 1966;61:455–460. doi: 10.1037/h0023263. [DOI] [PubMed] [Google Scholar]

[R182] 182.Li T, et al. Intranasal Oxytocin Restores Maternal Behavior and Oxytocin Neuronal Activity in the Supraoptic Nucleus in Rat Dams with Cesarean Delivery. Neuroscience. 2021;468:235–246. doi: 10.1016/j.neuroscience.2021.06.020. [DOI] [PubMed] [Google Scholar]

[R183] 183.Arnoux I, et al. Astrocytes control oxytocin-based maternal behavior via connexin 30. 2026 doi: 10.64898/2026.01.09.698569. Preprint at. [DOI] [Google Scholar]

[R184] 184.McNeilly AS, Robinson IC, Houston MJ, Howie PW. Release of oxytocin and prolactin in response to suckling. Br Med J. 1983;286:257–259. doi: 10.1136/bmj.286.6361.257. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R185] 185.Lucas BK, Ormandy CJ, Binart N, Bridges RS, Kelly PA. Null Mutation of the Prolactin Receptor Gene Produces a Defect in Maternal Behavior. Endocrinology. 1998;139 doi: 10.1210/endo.139.10.6243. [DOI] [PubMed] [Google Scholar]

[R186] 186.Yamaguchi T, Lin D. The hormonal and neural control of maternal aggression. Curr Opin Neurobiol. 2026;98:103194. doi: 10.1016/j.conb.2026.103194. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R187] 187.Stagkourakis S, et al. Maternal aggression driven by the transient mobilisation of a dormant hormone-sensitive circuit. Nat Commun. 2025;16:8553. doi: 10.1038/s41467-025-64043-4. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R188] 188.Yamaguchi T, et al. The neural mechanisms supporting the rise and fall of maternal aggression. Nature. 2026:1–11. doi: 10.1038/s41586-026-10354-5. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R189] 189.Georgescu T, Khant Aung Z, Grattan DR, Brown RSE. Prolactin-mediated restraint of maternal aggression in lactation. Proc Natl Acad Sci. 2022;119:e2116972119. doi: 10.1073/pnas.2116972119. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R190] 190.Shahrokh DK, Zhang T-Y, Diorio J, Gratton A, Meaney MJ. Oxytocin-Dopamine Interactions Mediate Variations in Maternal Behavior in the Rat. Endocrinology. 2010;151:2276–2286. doi: 10.1210/en.2009-1271. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R191] 191.Bosch OJ, Neumann ID. Brain vasopressin is an important regulator of maternal behavior independent of dams’ trait anxiety. Proc Natl Acad Sci. 2008;105:17139–17144. doi: 10.1073/pnas.0807412105. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R192] 192.Bosch OJ, Pförtsch J, Beiderbeck DI, Landgraf R, Neumann ID. Maternal Behaviour is Associated with Vasopressin Release in the Medial Preoptic Area and Bed Nucleus of the Stria Terminalis in the Rat. J Neuroendocrinol. 2010;22:420–429. doi: 10.1111/j.1365-2826.2010.01984.x. [DOI] [PubMed] [Google Scholar]

[R193] 193.Nephew B, Bridges R. Central actions of arginine vasopressin and a V1a receptor antagonist on maternal aggression, maternal behavior, and grooming in lactating rats. Pharmacol Biochem Behav. 2008;91:77–83. doi: 10.1016/j.pbb.2008.06.013. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R194] 194.Caughey SD, et al. Changes in the Intensity of Maternal Aggression and Central Oxytocin and Vasopressin V1a Receptors Across the Peripartum Period in the Rat. J Neuroendocrinol. 2011;23:1113–1124. doi: 10.1111/j.1365-2826.2011.02224.x. [DOI] [PubMed] [Google Scholar]

[R195] 195.Bayerl DS, Klampfl SM, Bosch OJ. Central V1b Receptor Antagonism in Lactating Rats: Impairment of Maternal Care But Not of Maternal Aggression. J Neuroendocrinol. 2014;26:918–926. doi: 10.1111/jne.12226. [DOI] [PubMed] [Google Scholar]

[R196] 196.Aspesi D, et al. The deletion of the arginine vasopressin 1a receptor impairs sexual and maternal behavior. Front Endocrinol. 2025;16:1649706. doi: 10.3389/fendo.2025.1649706. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R197] 197.Dvorkin R, Shea SD. Precise and Pervasive Phasic Bursting in Locus Coeruleus during Maternal Behavior in Mice. J Neurosci. 2022;42:2986–2999. doi: 10.1523/JNEUROSCI.0938-21.2022. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R198] 198.Corona A, Choe J, Muñoz-Castañeda R, Osten P, Shea SD. A circuit from the locus coeruleus to the anterior cingulate cortex modulates offspring interactions in mice. Cell Rep. 2023;42:112771. doi: 10.1016/j.celrep.2023.112771. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R199] 199.Holschbach MA, Vitale EM, Lonstein JS. Serotonin-specific lesions of the dorsal raphe disrupt maternal aggression and caregiving in postpartum rats. Behav Brain Res. 2018;348:53–64. doi: 10.1016/j.bbr.2018.04.008. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R200] 200.Muzerelle A, et al. Dorsal raphe serotonin neurotransmission is required for the expression of nursing behavior and for pup survival. Sci Rep. 2021;11:6004. doi: 10.1038/s41598-021-84368-6. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R201] 201.Gao J, Wu R, Davis C, Li M. Activation of 5-HT2A receptor disrupts rat maternal behavior. Neuropharmacology. 2018;128:96–105. doi: 10.1016/j.neuropharm.2017.09.037. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R202] 202.Hammer RP, Jr, Mateo AR, Bridges RS. Hormonal Regulation of Medial Preoptic μ-Opiate Receptor Density before and after Parturition. Neuroendocrinology. 1992;56:38–45. doi: 10.1159/000126206. [DOI] [PubMed] [Google Scholar]

[R203] 203.Mann PE, Pasternak GW, Bridges RS. Mu1 opioid receptor involvement in maternal behavior. Physiol Behav. 1990;47:133–138. doi: 10.1016/0031-9384(90)90051-5. [DOI] [PubMed] [Google Scholar]

[R204] 204.Byrnes EM, Rigero BA, Bridges RS. Opioid receptor antagonism during early lactation results in the increased duration of nursing bouts. Physiol Behav. 2000;70:211–216. doi: 10.1016/s0031-9384(00)00231-6. [DOI] [PubMed] [Google Scholar]

[R205] 205.Yip SH, et al. Elevated Prolactin during Pregnancy Drives a Phenotypic Switch in Mouse Hypothalamic Dopaminergic Neurons. Cell Rep. 2019;26:1787–1799.:e5. doi: 10.1016/j.celrep.2019.01.067. [DOI] [PubMed] [Google Scholar]

[R206] 206.Stolzenberg DS, Mayer HS. Experience-dependent mechanisms in the regulation of parental care. Front Neuroendocrinol. 2019;54:100745. doi: 10.1016/j.yfrne.2019.04.002. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R207] 207.Nephew BC, Bridges RS, Lovelock DF, Byrnes EM. Enhanced maternal aggression and associated changes in neuropeptide gene expression in multiparous rats. Behav Neurosci. 2009;123:949–957. doi: 10.1037/a0016734. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R208] 208.Ladyman SR, Khant Aung Z, Grattan DR. Impact of Pregnancy and Lactation on the Long-Term Regulation of Energy Balance in Female Mice. Endocrinology. 2018;159:2324–2336. doi: 10.1210/en.2018-00057. [DOI] [PubMed] [Google Scholar]

[R209] 209.Chechko N, Nehls S. From Pregnancy to Postpartum: The Dynamic Reorganization of the Maternal Brain. Neurosci Insights. 2025;20:26331055251315488. doi: 10.1177/26331055251315488. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R210] 210.Duarte-Guterman P, et al. Cellular and molecular signatures of motherhood in the adult and ageing rat brain. Open Biol. 2023;13:230217. doi: 10.1098/rsob.230217. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R211] 211.De Lange A-MG, et al. Population-based neuroimaging reveals traces of childbirth in the maternal brain. Proc Natl Acad Sci. 2019;116:22341–22346. doi: 10.1073/pnas.1910666116. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R212] 212.Pritschet L, et al. Neuroanatomical changes observed over the course of a human pregnancy. Nat Neurosci. 2024;27:2253–2260. doi: 10.1038/s41593-024-01741-0. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R213] 213.Brent LJN, et al. Ecological Knowledge, Leadership, and the Evolution of Menopause in Killer Whales. Curr Biol. 2015;25:746–750. doi: 10.1016/j.cub.2015.01.037. [DOI] [PubMed] [Google Scholar]

[R214] 214.Nattrass S, et al. Postreproductive killer whale grandmothers improve the survival of their grandoffspring. Proc Natl Acad Sci. 2019;116:26669–26673. doi: 10.1073/pnas.1903844116. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R215] 215.Lahdenperä M, Mar KU, Lummaa V. Nearby grandmother enhances calf survival and reproduction in Asian elephants. Sci Rep. 2016;6:27213. doi: 10.1038/srep27213. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R216] 216.Chapman SN, Lahdenperä M, Pettay JE, Lynch RF, Lummaa V. Offspring fertility and grandchild survival enhanced by maternal grandmothers in a pre-industrial human society. Sci Rep. 2021;11:3652. doi: 10.1038/s41598-021-83353-3. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R217] 217.Alcantara IC, et al. A hypothalamic circuit that modulates feeding and parenting behaviours. Nature. 2025;645:981–990. doi: 10.1038/s41586-025-09268-5. [DOI] [PubMed] [Google Scholar]

[R218] 218.Li X-Y, et al. AGRP Neurons Project to the Medial Preoptic Area and Modulate Maternal Nest-Building. J Neurosci. 2019;39:456–471. doi: 10.1523/JNEUROSCI.0958-18.2018. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R219] 219.Han Y, Li X-Y, Wang S-R, Wei Y-C, Xu X-H. Presence of pups suppresses hunger-induced feeding in virgin adult mice of both sexes. Neuroscience. 2017;362:228–238. doi: 10.1016/j.neuroscience.2017.08.047. [DOI] [PubMed] [Google Scholar]

[R220] 220.Muroi Y, Ishii T. Neuropeptide Y is crucial for nutritional state-dependent regulation of maternal behavior. Psychoneuroendocrinology. 2015;51:392–402. doi: 10.1016/j.psyneuen.2014.09.022. [DOI] [PubMed] [Google Scholar]

[R221] 221.Zoubovsky SP, et al. Chronic psychosocial stress during pregnancy affects maternal behavior and neuroendocrine function and modulates hypothalamic CRH and nuclear steroid receptor expression. Transl Psychiatry. 2020;10:6. doi: 10.1038/s41398-020-0704-2. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R222] 222.Smith JW, Seckl JR, Evans AT, Costall B, Smythe JW. Gestational stress induces post-partum depression-like behaviour and alters maternal care in rats. Psychoneuroendocrinology. 2004;29:227–244. doi: 10.1016/s0306-4530(03)00025-8. [DOI] [PubMed] [Google Scholar]

[R223] 223.Brummelte S, Galea LAM. Chronic corticosterone during pregnancy and postpartum affects maternal care, cell proliferation and depressive-like behavior in the dam. Horm Behav. 2010;58:769–779. doi: 10.1016/j.yhbeh.2010.07.012. [DOI] [PubMed] [Google Scholar]

[R224] 224.Pedersen CA, Caldwell JD, McGuire M, Evans DL. Corticotropin-releasing hormone inhibits maternal behavior and induces pup-killing. Life Sci. 1991;48:1537–1546. doi: 10.1016/0024-3205(91)90278-j. [DOI] [PubMed] [Google Scholar]

[R225] 225.Klampfl SM, et al. Maternal stress and the MPOA: Activation of CRF receptor 1 impairs maternal behavior and triggers local oxytocin release in lactating rats. Neuropharmacology. 2018;133:440–450. doi: 10.1016/j.neuropharm.2018.02.019. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R226] 226.Klampfl SM, et al. Brain CRF-binding protein modulates aspects of maternal behavior under stressful conditions and supports a hypo-anxious state in lactating rats. Horm Behav. 2016;84:136–144. doi: 10.1016/j.yhbeh.2016.06.009. [DOI] [PubMed] [Google Scholar]

[R227] 227.Rivero-Quiles C, et al. Appetitive maternal behavior in mice is resistant to perinatal chronic variable stress and perinatal chronic corticosterone treatment. Horm Behav. 2026;177:105855. doi: 10.1016/j.yhbeh.2025.105855. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R228] 228.Erskine MS. Solicitation behavior in the estrous female rat: a review. Horm Behav. 1989;23:473–502. doi: 10.1016/0018-506x(89)90037-8. [DOI] [PubMed] [Google Scholar]

[R229] 229.Pfaff DW, Sakuma Y. Deficit in the lordosis reflex of female rats caused by lesions in the ventromedial nucleus of the hypothalamus. J Physiol. 1979;288:203–210. [PMC free article] [PubMed] [Google Scholar]

[R230] 230.Gutierrez-Castellanos N, et al. A hypothalamic node for the cyclical control of female sexual rejection. Neuron. 2024:S0896-6273(24)00774–8. doi: 10.1016/j.neuron.2024.10.026. [DOI] [PubMed] [Google Scholar]

[R231] 231.Inoue S, et al. Periodic Remodeling in a Neural Circuit Governs Timing of Female Sexual Behavior. Cell. 2019;179:1393–1408.:e16. doi: 10.1016/j.cell.2019.10.025. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R232] 232.Yin L, et al. VMHvllCckar cells dynamically control female sexual behaviors over the reproductive cycle. Neuron. 2022;110:3000–3017.:e8. doi: 10.1016/j.neuron.2022.06.026. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R233] 233.Ishii KK, et al. Post-ejaculatory Inhibition of Female Sexual Drive via Heterogeneous Neuronal Ensembles in the Medial Preoptic Area. eLife. 2025;12 doi: 10.7554/eLife.91765. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R234] 234.Sun F, et al. Shared neural substrates of prosocial and parenting behaviours. Nature. 2026 doi: 10.1038/s41586-026-10327-8. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R235] 235.de Waal FBM. Putting the altruism back into altruism: the evolution of empathy. Annu Rev Psychol. 2008;59:279–300. doi: 10.1146/annurev.psych.59.103006.093625. [DOI] [PubMed] [Google Scholar]

[R236] 236.Niepoth N, et al. Evolution of a novel adrenal cell type that promotes parental care. Nature. 2024;629:1082–1090. doi: 10.1038/s41586-024-07423-y. [ This study discovers that monogamous oldfield mice recently evolved a novel adrenal gland cell type which promotes caregiving behaviours by converting progesterone into 20α-hydroxyprogesterone ] [DOI] [PMC free article] [PubMed] [Google Scholar]

[R237] 237.Young KA, Gobrogge KL, Liu Y, Wang Z. The neurobiology of pair bonding: insights from a socially monogamous rodent. Front Neuroendocrinol. 2011;32:53–69. doi: 10.1016/j.yfrne.2010.07.006. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R238] 238.Rogers FD, et al. Agouti integrates environmental cues to regulate paternal behaviour. Nature. 2026;652:694–702. doi: 10.1038/s41586-026-10123-4. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R239] 239.Li Y, et al. Paternal behavior is controlled by preoptic Trpc5 neurons. 2026 doi: 10.64898/2026.03.20.713203. Preprint at. [DOI] [Google Scholar]

[R240] 240.Vom Saal FS, Howard LS. The Regulation of Infanticide and Parental Behavior: Implications for Reproductive Success in Male Mice. Science. 1982;215:1270–1272. doi: 10.1126/science.7058349. [DOI] [PubMed] [Google Scholar]

[R241] 241.Vom Saal FS. Time-contingent change in infanticide and parental behavior induced by ejaculation in male mice. Physiol Behav. 1985;34:7–15. doi: 10.1016/0031-9384(85)90069-1. [DOI] [PubMed] [Google Scholar]

[R242] 242.McCarthy MM, Vom Saal FS. Inhibition of infanticide after mating by wild male house mice. Physiol Behav. 1986;36:203–209. doi: 10.1016/0031-9384(86)90004-1. [DOI] [PubMed] [Google Scholar]

[R243] 243.Perrigo G, Bryant WC, Vom Saal FS. A unique neural timing system prevents male mice from harming their own offspring. Anim Behav. 1990;39:535–539. [Google Scholar]

[R244] 244.Rosenfeld CS, Johnson SA, Ellersieck MR, Roberts RM. Interactions between Parents and Parents and Pups in the Monogamous California Mouse (Peromyscus californicus) PLoS ONE. 2013;8:e75725. doi: 10.1371/journal.pone.0075725. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R245] 245.Valera-Marín G, et al. Raised without a father: monoparental care effects over development, sexual behavior, sexual reward, and pair bonding in prairie voles. Behav Brain Res. 2021;408:113264. doi: 10.1016/j.bbr.2021.113264. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R246] 246.Clutton-Brock TH, et al. Effects of Helpers on Juvenile Development and Survival in Meerkats. Science. 2001;293:2446–2449. doi: 10.1126/science.1061274. [DOI] [PubMed] [Google Scholar]

[R247] 247.Ziegler TE, Prudom SL, Zahed SR. Variations in male parenting behavior and physiology in the common marmoset. Am J Hum Biol. 2009;21:739–744. doi: 10.1002/ajhb.20920. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R248] 248.Lukas D, Clutton-Brock T. Life histories and the evolution of cooperative breeding in mammals. Proc R Soc B Biol Sci. 2012;279:4065–4070. doi: 10.1098/rspb.2012.1433. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R249] 249.Russell AF, Young AJ, Spong G, Jordan NR, Clutton-Brock TH. Helpers increase the reproductive potential of offspring in cooperative meerkats. Proc R Soc B Biol Sci. 2007;274:513–520. doi: 10.1098/rspb.2006.3698. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R250] 250.Rosenbaum S, Gettler LT. With a little help from her friends (and family) part I: the ecology and evolution of non-maternal care in mammals. Physiol Behav. 2018;193:1–11. doi: 10.1016/j.physbeh.2017.12.025. [DOI] [PubMed] [Google Scholar]

[R251] 251.Kentner AC, Abizaid A, Bielajew C. Modeling Dad: Animal models of paternal behavior. Neurosci Biobehav Rev. 2010;34:438–451. doi: 10.1016/j.neubiorev.2009.08.010. [DOI] [PubMed] [Google Scholar]

[R252] 252.Gubernick DJ, Alberts JR. The biparental care system of the California mouse, Peromyscus californicus. J Comp Psychol. 1987;101:169–177. [PubMed] [Google Scholar]

[R253] 253.Ahern TH, Hammock EAD, Young LJ. Parental division of labor, coordination, and the effects of family structure on parenting in monogamous prairie voles (Microtus ochrogaster) Dev Psychobiol. 2011;53:118–131. doi: 10.1002/dev.20498. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R254] 254.Bendesky A, et al. The genetic basis of parental care evolution in monogamous mice. Nature. 2017;544:434–439. doi: 10.1038/nature22074. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R255] 255.Khadraoui M, Merritt JR, Hoekstra HE, Bendesky A. Post-mating parental behavior trajectories differ across four species of deer mice. PLOS ONE. 2022;17:e0276052. doi: 10.1371/journal.pone.0276052. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R256] 256.Elwood RW. Changes in the responses of male and female gerbils (Meriones unguiculatus) towards test pups during the pregnancy of the female. Anim Behav. 1977;25:46–51. [Google Scholar]

[R257] 257.Elwood RW, Stolzenberg DS. Flipping the parental switch: from killing to caring in male mammals. Anim Behav. 2020;165:133–142. doi: 10.1016/j.anbehav.2020.05.001. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R258] 258.Kennedy HF, Elwood RW. Strain differences in the inhibition of infanticide in male mice (Mus musculus) Behav Neural Biol. 1988;50:349–353. doi: 10.1016/s0163-1047(88)91066-7. [DOI] [PubMed] [Google Scholar]

[R259] 259.Palanza P, Parmigiani S. Inhibition of infanticide in male Swiss mice: Behavioral polymorphism in response to multiple mediating factors. Physiol Behav. 1991;49:797–802. doi: 10.1016/0031-9384(91)90320-n. [DOI] [PubMed] [Google Scholar]

[R260] 260.Lee AW, Brown RE. Medial preoptic lesions disrupt parental behavior in both male and female California mice (Peromyscus californicus) Behav Neurosci. 2002;116:968–975. doi: 10.1037//0735-7044.116.6.968. [DOI] [PubMed] [Google Scholar]

[R261] 261.Lee AW, Brown RE. Comparison of medial preoptic, amygdala, and nucleus accumbens lesions on parental behavior in California mice (Peromyscus californicus) Physiol Behav. 2007;92:617–628. doi: 10.1016/j.physbeh.2007.05.008. [DOI] [PubMed] [Google Scholar]

[R262] 262.Ito K, et al. Synaptic plasticity in the medial preoptic area of male mice encodes social experiences with female and regulates behavior toward young. eLife. 2025;14 [ This study proposes a two-stage model in which distinct forms of synaptic plasticity in the MPOA sequentially suppress infanticide and promote the emergence of paternal care after mating in mice ] [Google Scholar]

[R263] 263.Inada K, et al. Vasopressin-to-oxytocin receptor crosstalk in the preoptic area underlying parental behaviors in male mice. Nat Commun. 2025;16:10844. doi: 10.1038/s41467-025-66908-0. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R264] 264.He Z, et al. Paraventricular Nucleus Oxytocin Subsystems Promote Active Paternal Behaviors in Mandarin Voles. J Neurosci Off J Soc Neurosci. 2021;41:6699–6713. doi: 10.1523/JNEUROSCI.2864-20.2021. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R265] 265.Trainor BC, Marler CA. Testosterone, Paternal Behavior, and Aggression in the Monogamous California Mouse (Peromyscus californicus) Horm Behav. 2001;40:32–42. doi: 10.1006/hbeh.2001.1652. [DOI] [PubMed] [Google Scholar]

[R266] 266.Trainor BC, Bird IM, Alday NA, Schlinger BA, Marler CA. Variation in Aromatase Activity in the Medial Preoptic Area and Plasma Progesterone Is Associated with the Onset of Paternal Behavior. Neuroendocrinology. 2003;78:36–44. doi: 10.1159/000071704. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R267] 267.Gleason ED, Marler CA. Testosterone response to courtship predicts future paternal behavior in the California mouse, Peromyscus californicus. Horm Behav. 2010;57:147–154. doi: 10.1016/j.yhbeh.2009.10.006. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R268] 268.Wang Z, De Vries GJ. Testosterone effects on paternal behavior and vasopressin immunoreactive projections in prairie voles (Microtus ochrogaster) Brain Res. 1993;631:156–160. doi: 10.1016/0006-8993(93)91203-5. [DOI] [PubMed] [Google Scholar]

[R269] 269.Lonstein, De Vries Sex Differences in the Parental Behaviour of Adult Virgin Prairie Voles: Independence From Gonadal Hormones and Vasopressin. J Neuroendocrinol. 1999;11:441–449. doi: 10.1046/j.1365-2826.1999.00361.x. [DOI] [PubMed] [Google Scholar]

[R270] 270.Hume JM, Wynne-Edwards KE. Castration reduces male testosterone, estradiol, and territorial aggression, but not paternal behavior in biparental dwarf hamsters (Phodopus campbelli) Horm Behav. 2005;48:303–310. doi: 10.1016/j.yhbeh.2005.04.001. [DOI] [PubMed] [Google Scholar]

[R271] 271.Saxbe DE, et al. Fathers’ decline in testosterone and synchrony with partner testosterone during pregnancy predicts greater postpartum relationship investment. Horm Behav. 2017;90:39–47. doi: 10.1016/j.yhbeh.2016.07.005. [DOI] [PubMed] [Google Scholar]

[R272] 272.Meijer WM, Van IJzendoorn MH, Bakermans-Kranenburg MJ. Challenging the challenge hypothesis on testosterone in fathers: Limited meta-analytic support. Psychoneuroendocrinology. 2019;110:104435. doi: 10.1016/j.psyneuen.2019.104435. [DOI] [PubMed] [Google Scholar]

[R273] 273.Gettler LT, McDade TW, Feranil AB, Kuzawa CW. Longitudinal evidence that fatherhood decreases testosterone in human males. Proc Natl Acad Sci. 2011;108:16194–16199. doi: 10.1073/pnas.1105403108. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R274] 274.Delahunty KM, McKay DW, Noseworthy DE, Storey AE. Prolactin responses to infant cues in men and women: Effects of parental experience and recent infant contact. Horm Behav. 2007;51:213–220. doi: 10.1016/j.yhbeh.2006.10.004. [DOI] [PubMed] [Google Scholar]

[R275] 275.Gordon I, Zagoory-Sharon O, Leckman JF, Feldman R. Prolactin, Oxytocin, and the development of paternal behavior across the first six months of fatherhood. Horm Behav. 2010;58:513–518. doi: 10.1016/j.yhbeh.2010.04.007. [DOI] [PMC free article] [PubMed] [Google Scholar]

[R276] 276.Feldman R, Gordon I, Schneiderman I, Weisman O, Zagoory-Sharon O. Natural variations in maternal and paternal care are associated with systematic changes in oxytocin following parent–infant contact. Psychoneuroendocrinology. 2010;35:1133–1141. doi: 10.1016/j.psyneuen.2010.01.013. [DOI] [PubMed] [Google Scholar]

PERMALINK

Neural circuits for mammalian parental behaviour

Bradley B Jamieson

Johannes Kohl

Abstract

Introduction

Diversity of mammalian parenting

Core parental circuitry

Fig. 1. Functional circuit logic underlying rodent infant-directed behaviours.

Evaluating infant cues

Pro- and anti-parental central systems

Reinforcement and action

Cortical contributions

Parental behaviour across the lifespan

Fig. 2. Parental behaviour across the lifespan.

Onset of alloparental behaviour in juveniles

Fig. 3. Caregiving outside parenthood.

Learning to care: experience-driven parenting

Reproductive experience and parenthood

Fig. 4. Hormone action in parenting circuits during motherhood.

Pregnancy

Parturition and parenthood

Long-term plasticity and parental experience

Balancing care with competing drives

Conclusions and perspectives

Box 1. Mammalian parenting strategies.

Box 2. Paternal care.

Acknowledgements

Glossary

Footnotes

References

ACTIONS

PERMALINK

RESOURCES

Cite

Add to Collections

PERMALINK

Neural circuits for mammalian parental behaviour

Bradley B Jamieson

Johannes Kohl

Abstract

Introduction

Diversity of mammalian parenting

Core parental circuitry

Fig. 1. Functional circuit logic underlying rodent infant-directed behaviours.

Evaluating infant cues

Pro- and anti-parental central systems

Reinforcement and action

Cortical contributions

Parental behaviour across the lifespan

Fig. 2. Parental behaviour across the lifespan.

Onset of alloparental behaviour in juveniles

Fig. 3. Caregiving outside parenthood.

Learning to care: experience-driven parenting

Reproductive experience and parenthood

Fig. 4. Hormone action in parenting circuits during motherhood.

Pregnancy

Parturition and parenthood

Long-term plasticity and parental experience

Balancing care with competing drives

Conclusions and perspectives

Box 1. Mammalian parenting strategies.

Box 2. Paternal care.

Acknowledgements

Glossary

Footnotes

References

ACTIONS

PERMALINK

RESOURCES

Similar articles

Cited by other articles

Links to NCBI Databases