Prentice, A. M. & Prentice, A. Energy costs of lactation. Annu. Rev. Nutr. 8, 63–79 (1988).
Fleming, A. S. & Rosenblatt, J. S. Maternal behavior in the virgin and lactating rat. J. Comp. Physiol. Psychol. 86, 957–972 (1974).
Brunton, P. J. & Russell, J. A. The expectant brain: adapting for motherhood. Nat. Rev. Neurosci. 9, 11–25 (2008).
Numan, M. & Woodside, B. Maternity: neural mechanisms, motivational processes, and physiological adaptations. Behav. Neurosci. 124, 715–741 (2010).
Woodside, B., Budin, R., Wellman, M. K. & Abizaid, A. Many mouths to feed: the control of food intake during lactation. Front. Neuroendocrinol. 33, 301–314 (2012).
Sohn, J. W., Elmquist, J. K. & Williams, K. W. Neuronal circuits that regulate feeding behavior and metabolism. Trends Neurosci. 36, 504–512 (2013).
Ahn, B. H., Kim, M. & Kim, S. Y. Brain circuits for promoting homeostatic and non-homeostatic appetites. Exp. Mol. Med. 54, 349–357 (2022).
Alcantara, I. C., Tapia, A. P. M., Aponte, Y. & Krashes, M. J. Acts of appetite: neural circuits governing the appetitive, consummatory, and terminating phases of feeding. Nat. Metab. 4, 836–847 (2022).
Kohl, J. Parenting—a paradigm for investigating the neural circuit basis of behavior. Curr. Opin. Neurobiol. 60, 84–91 (2020).
Burnett, C. J. et al. Hunger-driven motivational state competition. Neuron 92, 187–201 (2016).
Burnett, C. J. et al. Need-based prioritization of behavior. eLife 8, e44527 (2019).
Sutton, A. K. & Krashes, M. J. Integrating hunger with rival motivations. Trends Endocrinol. Metab. 31, 495–507 (2020).
Barajas-Azpeleta, R., Tastekin, I. & Ribeiro, C. Neuroscience: How the brain prioritizes behaviors. Curr. Biol. 31, R1125–R1127 (2021).
Kovac, M. P. & Davis, W. J. Neural mechanism underlying behavioral choice in Pleurobranchaea. J. Neurophysiol. 43, 469–487 (1980).
Hong, W., Kim, D. W. & Anderson, D. J. Antagonistic control of social versus repetitive self-grooming behaviors by separable amygdala neuronal subsets. Cell 158, 1348–1361 (2014).
Cheriyamkunnel, S. J. et al. A neuronal mechanism controlling the choice between feeding and sexual behaviors in Drosophila. Curr. Biol. 31, 4231–4245.e4 (2021).
Mei, L., Yan, R., Yin, L., Sullivan, R. M. & Lin, D. Antagonistic circuits mediating infanticide and maternal care in female mice. Nature 618, 1006–1016 (2023).
de Araujo Salgado, I. et al. Toggling between food-seeking and self-preservation behaviors via hypothalamic response networks. Neuron 111, 2899–2917.e6 (2023).
Marlin, B. J., Mitre, M., D’Amour J, A., Chao, M. V. & Froemke, R. C. Oxytocin enables maternal behaviour by balancing cortical inhibition. Nature 520, 499–504 (2015).
Stolzenberg, D. S. & Champagne, F. A. Hormonal and non-hormonal bases of maternal behavior: The role of experience and epigenetic mechanisms. Horm. Behav. 77, 204–210 (2016).
Ammari, R. et al. Hormone-mediated neural remodeling orchestrates parenting onset during pregnancy. Science 382, 76–81 (2023).
Matikainen-Ankney, B. A. et al. An open-source device for measuring food intake and operant behavior in rodent home-cages. eLife 10, e66173 (2021).
Campbell, J. N. et al. A molecular census of arcuate hypothalamus and median eminence cell types. Nat. Neurosci. 20, 484–496 (2017).
Krashes, M. J. et al. Rapid, reversible activation of AgRP neurons drives feeding behavior in mice. J. Clin. Invest. 121, 1424–1428 (2011).
Aponte, Y., Atasoy, D. & Sternson, S. M. AGRP neurons are sufficient to orchestrate feeding behavior rapidly and without training. Nat. Neurosci. 14, 351–355 (2011).
Takahashi, K. A. & Cone, R. D. Fasting induces a large, leptin-dependent increase in the intrinsic action potential frequency of orexigenic arcuate nucleus neuropeptide Y/Agouti-related protein neurons. Endocrinology 146, 1043–1047 (2005).
Mandelblat-Cerf, Y. et al. Arcuate hypothalamic AgRP and putative POMC neurons show opposite changes in spiking across multiple timescales. eLife 4, e07122 (2015).
Chen, Y., Lin, Y. C., Kuo, T. W. & Knight, Z. A. Sensory detection of food rapidly modulates arcuate feeding circuits. Cell 160, 829–841 (2015).
Betley, J. N. et al. Neurons for hunger and thirst transmit a negative-valence teaching signal. Nature 521, 180–185 (2015).
Padilla, S. L. et al. Agouti-related peptide neural circuits mediate adaptive behaviors in the starved state. Nat. Neurosci. 19, 734–741 (2016).
Alhadeff, A. L. et al. A Neural circuit for the suppression of pain by a competing need state. Cell 173, 140–152.e15 (2018).
Li, X. Y. et al. AGRP neurons project to the medial preoptic area and modulate maternal nest-building. J. Neurosci. 39, 456–471 (2019).
Numan, M. Medial preoptic area and maternal behavior in the female rat. J. Comp. Physiol. Psychol. 87, 746–759 (1974).
Numan, M., Numan, M. J., Marzella, S. R. & Palumbo, A. Expression of c-fos, fos B, and egr-1 in the medial preoptic area and bed nucleus of the stria terminalis during maternal behavior in rats. Brain Res. 792, 348–352 (1998).
Wu, Z., Autry, A. E., Bergan, J. F., Watabe-Uchida, M. & Dulac, C. G. Galanin neurons in the medial preoptic area govern parental behaviour. Nature 509, 325–330 (2014).
Kohl, J. et al. Functional circuit architecture underlying parental behaviour. Nature 556, 326–331 (2018).
Fang, Y. Y., Yamaguchi, T., Song, S. C., Tritsch, N. X. & Lin, D. A hypothalamic midbrain pathway essential for driving maternal behaviors. Neuron 98, 192–207.e10 (2018).
Brown, R. S. E. et al. Prolactin action in the medial preoptic area is necessary for postpartum maternal nursing behavior. Proc. Natl Acad. Sci. USA 114, 10779–10784 (2017).
Yoshihara, C. et al. Calcitonin receptor signaling in the medial preoptic area enables risk-taking maternal care. Cell Rep. 35, 109204 (2021).
Atasoy, D., Betley, J. N., Su, H. H. & Sternson, S. M. Deconstruction of a neural circuit for hunger. Nature 488, 172–177 (2012).
Betley, J. N., Cao, Z. F., Ritola, K. D. & Sternson, S. M. Parallel, redundant circuit organization for homeostatic control of feeding behavior. Cell 155, 1337–1350 (2013).
Moffitt, J. R. et al. Molecular, spatial, and functional single-cell profiling of the hypothalamic preoptic region. Science 362, eaau5324 (2018).
Guenthner, C. J., Miyamichi, K., Yang, H. H., Heller, H. C. & Luo, L. Permanent genetic access to transiently active neurons via TRAP: targeted recombination in active populations. Neuron 78, 773–784 (2013).
DeNardo, L. A. et al. Temporal evolution of cortical ensembles promoting remote memory retrieval. Nat. Neurosci. 22, 460–469 (2019).
Cazzulino, A. S., Martinez, R., Tomm, N. K. & Denny, C. A. Improved specificity of hippocampal memory trace labeling. Hippocampus 26, 752–762 (2016).
McHenry, J. A. et al. Hormonal gain control of a medial preoptic area social reward circuit. Nat. Neurosci. 20, 449–458 (2017).
Ohki-Hamazaki, H. et al. Mice lacking bombesin receptor subtype-3 develop metabolic defects and obesity. Nature 390, 165–169 (1997).
Guan, X. M. et al. Antiobesity effect of MK-5046, a novel bombesin receptor subtype-3 agonist. J. Pharmacol. Exp. Ther. 336, 356–364 (2011).
Pinol, R. A. et al. Preoptic BRS3 neurons increase body temperature and heart rate via multiple pathways. Cell Metab. 33, 1389–1403.e6 (2021).
Motta, S. C. et al. Ventral premammillary nucleus as a critical sensory relay to the maternal aggression network. Proc. Natl Acad. Sci. USA 110, 14438–14443 (2013).
Donato, J. Jr. et al. Lesions of the ventral premammillary nucleus disrupt the dynamic changes in Kiss1 and GnRH expression characteristic of the proestrus-estrus transition. Neuroscience 241, 67–79 (2013).
Donato, J. Jr. et al. The ventral premammillary nucleus links fasting-induced changes in leptin levels and coordinated luteinizing hormone secretion. J. Neurosci. 29, 5240–5250 (2009).
Mei, L., Osakada, T. & Lin, D. Hypothalamic control of innate social behaviors. Science 382, 399–404 (2023).
Chen, P., Li, C., Haskell-Luevano, C., Cone, R. D. & Smith, M. S. Altered expression of agouti-related protein and its colocalization with neuropeptide Y in the arcuate nucleus of the hypothalamus during lactation. Endocrinology 140, 2645–2650 (1999).
Phillips, C. T. & Palmiter, R. D. Role of agouti-related protein-expressing neurons in lactation. Endocrinology 149, 544–550 (2008).
Suzuki, Y. et al. Changes in mRNA expression of arcuate nucleus appetite-regulating peptides during lactation in rats. J. Mol. Endocrinol. 52, 97–109 (2014).
Catalbas, K. et al. Hypothalamic AgRP neurons regulate the hyperphagia of lactation. Mol. Metab. 86, 101975 (2024).
Szymczak-Workman, A. L., Vignali, K. M. & Vignali, D. A. Verification of 2 A peptide cleavage. Cold Spring Harb. Protoc. 2012, 255–257 (2012).
Raymond, C. S. & Soriano, P. High-efficiency FLP and PhiC31 site-specific recombination in mammalian cells. PLoS ONE 2, e162 (2007).
Mickelsen, L. E. et al. Neurochemical heterogeneity among lateral hypothalamic hypocretin/orexin and melanin-concentrating hormone neurons identified through single-cell gene expression analysis. eNeuro https://doi.org/10.1523/ENEURO.0013-17.2017 (2017).
Mickelsen, L. E. et al. Cellular taxonomy and spatial organization of the murine ventral posterior hypothalamus. eLife 9, e58901 (2020).
Zheng, G. X. et al. Massively parallel digital transcriptional profiling of single cells. Nat. Commun. 8, 14049 (2017).
Hao, Y. et al. Integrated analysis of multimodal single-cell data. Cell 184, 3573–3587.e29 (2021).
Saunders, A. et al. Molecular diversity and specializations among the cells of the adult mouse brain. Cell 174, 1015–1030.e16 (2018).
Friard, O. & Gamba, M. BORIS: a free, versatile open-source event-logging software for video/audio coding and live observations. Methods Ecol. Evol. 7, 1325–1330 (2016).
Gao, C. et al. Molecular and spatial profiling of the paraventricular nucleus of the thalamus. eLife 12, e81818 (2023).
Stirling, D. R. et al. CellProfiler 4: improvements in speed, utility and usability. BMC Bioinformatics 22, 433 (2021).
Alcantara, I. C., Papas, B. N., & Krashes, M. J. Single-cell RNA-sequencing of the ARC and MPOA across satiety and lactation states in female mice. Zenodo https://doi.org/10.5281/zenodo.15319501 (2025).
