Vasopressin and oxytocin release within the brain: a dynamic concept of multiple and variable modes of neuropeptide communication

Landgraf, Rainer; Neumann, Inga D.

doi:10.1016/j.yfrne.2004.05.001

Cited by 1,059Neuroendocrine regulation and behaviorNeuropeptides and Animal PhysiologyNeuroscience and Neuropharmacology Research

Vasopressin and oxytocin release within the brain: a dynamic concept of multiple and variable modes of neuropeptide communication

Landgraf, Rainer, Neumann, Inga D.·2004·Frontiers in Neuroendocrinology

DOI 10.1016/j.yfrne.2004.05.001

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Abstract

As exemplified particularly with vasopressin and oxytocin, release of neuropeptides within the brain occurs from dendrites, somata, and axons of neurosecretory neurons; mechanisms include activation of intracellular Ca2+ stores, changed strength of synaptic input and altered interaction between transcription factors and gene promoters. Upon demand, both diffuse spread of neuropeptides in the extracellular fluid following dendritic release and focal release from axonal terminals may contribute to regionally and temporally varying combinations of neuromodulator and neurotransmitter actions, thus providing a theoretically unlimited variability in interneuronal signaling. Thus, instead of favoring volume or synaptic transmission following central neuropeptide release, a more dynamic concept is presented with multiple and variable modes of release and communication. This concept considers neuropeptides in the extracellular fluid of the brain rather than those in the cerebrospinal fluid or plasma as primary signals, triggering a variety of receptor-mediated effects, including those underlying behavioral and neuroendocrine regulation and psychopathology.

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Effects of a single neuropharmacological challenge might differ considerably from endogenous OT mechanisms.
“effects of a single neuropharmacological challenge might differ considerably from endogenous OT mechanisms”
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The PVN and SON are the primary producers of OXT, which is released into both the blood and the brain in response to reproductive stimuli (e.g. birth, suckling, sex) as well as social interaction and stressors.
“These regions are the primary producers of OXT, which is released into both the blood and the brain in response to reproductive stimuli (e.g. birth, suckling, sex) as well as social interaction and stressors (Landgraf and Neumann, 2004).”
From Convergence of oxytocin and dopamine signalling in neuronal circuits: Insights into the neurobiology of social interactions across species (2024) · cites this paper as Landgraf and Neumann, 2004
A more global form of OT and AVP release can occur when these neuropeptides are released from non-synaptic regions of neurons, such as dendrites.
“A more global form of OT and AVP release can occur when these neuropeptides are released from non-synaptic regions of neurons (e.g., dendrites) (Figure 6) [108; 109; 110].”
From Cross-talk among oxytocin and arginine-vasopressin receptors: Relevance for basic and clinical studies of the brain and periphery (2018) · cites this paper as [109]
Published case studies and anecdotal evidence suggest off-label prescriptions of oxytocin are already a common practice by treating clinicians, despite concerns about pediatric use outside of a clinical trial setting.
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From Beyond the hype and hope: critical considerations for intranasal oxytocin research in autism spectrum disorder (2017) · cites this paper as [69]

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Stimulus-Dependent Translocation of κ Opioid Receptors to the Plasma Membrane
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RNA, Whither Goest Thou?
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Molecular components of the exocytotic machinery in the rat pituitary gland.
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Electrical stimulations of perifused magnocellular nuclei in vitro elicit Ca2+-dependent, tetrodotoxin-insensitive release of oxytocin and vasopressin
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Vasopressin released within the septal brain area during swim stress modulates the behavioural stress response in rats
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Forced swimming stimulates the expression of vasopressin and oxytocin in magnocellular neurons of the rat hypothalamic paraventricular nucleus
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The action of oxytocin originating in the hypothalamic paraventricular nucleus on mitral and granule cells in the rat main olfactory bulb
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Vasopressin facilitates its own release within the rat supraoptic nucleus in vivo
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Peptides and blood-brain barrier transport
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Endogenous opioid regulation of stress-induced oxytocin release within the hypothalamic paraventricular nucleus is reversed in late pregnancy: a microdialysis study
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Vasopressin Differentially Modulates Non-NMDA Receptors in Vasopressin and Oxytocin Neurons in the Supraoptic Nucleus
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Neurohormones in Invertebrates
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Oxytocin increases trust in humans
Kosfeld, M., Heinrichs, M., Zak, P. J., Fischbacher, U., & Fehr, E. (2005) · Nature · doi:10.1038/nature03701 · Cited by 3,690 · see citing works
Oxytocin and vasopressin in the human brain: social neuropeptides for translational medicine
Meyer-Lindenberg, A., Domes, G., Kirsch, P., & Heinrichs, M. (2011) · Nature Reviews Neuroscience · doi:10.1038/nrn3044 · Cited by 1,688 · see citing works
The neuro-symphony of stress
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Oxytocin improves “mind-reading” in humans
Domes, G., Heinrichs, M., Michel, A., Berger, C., & Herpertz, S. C. (2007) · Biological Psychiatry · doi:10.1016/j.biopsych.2006.07.015 · Cited by 1,298 · see citing works
Oxytocin shapes the neural circuitry of trust and trust adaptation in humans
Baumgartner, T., Heinrichs, M., Vonlanthen, A., Fischbacher, U., & Fehr, E. (2008) · Neuron · doi:10.1016/j.neuron.2008.04.009 · Cited by 1,258 · see citing works
Evoked axonal oxytocin release in the central amygdala attenuates fear response
Knobloch, H. S., Charlet, A., Hoffmann, L. C., Eliava, M., Khrulev, S., Cetin, A. H., Osten, P., Schwarz, M. K., Seeburg, P. H., Stoop, R., et al. (2012) · Neuron · doi:10.1016/j.neuron.2011.11.030 · Cited by 1,049 · see citing works
The oxytocin receptor: from intracellular signaling to behavior
Jurek, B., & Neumann, I. D. (2018) · Physiological Reviews · doi:10.1152/physrev.00031.2017 · Cited by 999 · see citing works
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Balance of brain oxytocin and vasopressin: implications for anxiety, depression, and social behaviors
Neumann, I. D., & Landgraf, R. (2012) · Trends in Neurosciences · doi:10.1016/j.tins.2012.08.004 · Cited by 954 · see citing works
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Dendritic peptide release and peptide-dependent behaviours
Ludwig, M., & Leng, G. (2006) · Nature Reviews Neuroscience · doi:10.1038/nrn1845 · Cited by 945 · see citing works
Promoting social behavior with oxytocin in high-functioning autism spectrum disorders.
Andari, E., Duhamel, J.-R., Zalla, T., Herbrecht, E., Leboyer, M., & Sirigu, A. (2010) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0910249107 · Cited by 941 · see citing works
Psychosocial and psychophysiological effects of human-animal interactions: the possible role of oxytocin
Beetz, A., Uvnäs-Moberg, K., Julius, H., & Kotrschal, K. (2012) · Frontiers in Psychology · doi:10.3389/fpsyg.2012.00234 · Cited by 904 · see citing works
“…in the hypothalamus and released into the circulatory system and the brain in response to sensory stimulation via a network of OT-containing nerves (Landgraf and Neumann, 2004; Ross et al., 2009), e.g., during breastfeeding, labor, sex, but also touch, warmth, and stroking, usually in the context…”
Oxytocin and the Neural Mechanisms Regulating Social Cognition and Affiliative Behavior
Ross, H. E., & Young, L. J. (2009) · Frontiers in Neuroendocrinology · doi:10.1016/j.yfrne.2009.05.004 · Cited by 856 · see citing works
“These multiple modes of central release of OT may allow high temporal and spatial resolution and a theoretically unlimited variability in OT signaling modes [119].”
BackgroundInfluential
Vasopressin and Oxytocin Excite Distinct Neuronal Populations in the Central Amygdala
Huber, D., Veinante, P., & Stoop, R. (2005) · Science · doi:10.1126/science.1105636 · Cited by 804 · see citing works
Oxytocin, vasopressin, and human social behavior
Heinrichs, M., von Dawans, B., & Domes, G. (2009) · Frontiers in Neuroendocrinology · doi:10.1016/j.yfrne.2009.05.005 · Cited by 799 · see citing works
Oxytocin attenuates amygdala responses to emotional faces regardless of valence
Domes, G., Heinrichs, M., Gläscher, J., Büchel, C., Braus, D. F., & Herpertz, S. C. (2007) · Biological Psychiatry · doi:10.1016/j.biopsych.2007.03.025 · Cited by 774 · see citing works
Oxytocin and social affiliation in humans
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Brain oxytocin: A key regulator of emotional and social behaviours in both females and males
Neumann, I. D. (2008) · Journal of Neuroendocrinology · doi:10.1111/j.1365-2826.2008.01726.x · Cited by 743 · see citing works
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Psychobiological mechanisms underlying the social buffering of the hypothalamic-pituitary-adrenocortical axis: a review of animal models and human studies across development
Hostinar, C. E., Sullivan, R. M., & Gunnar, M. R. (2014) · Psychological Bulletin · doi:10.1037/a0032671 · Cited by 739 · see citing works
The varieties of self-transcendent experience
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The Peptide That Binds: A Systematic Review of Oxytocin and its Prosocial Effects in Humans
MacDonald, K., & MacDonald, T. M. (2010) · Harvard Review of Psychiatry · doi:10.3109/10673220903523615 · Cited by 585 · see citing works
Early experience in humans is associated with changes in neuropeptides critical for regulating social behavior
Fries, A. B. W., Ziegler, T. E., Kurian, J. R., Jacoris, S., & Pollak, S. D. (2005) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0504767102 · Cited by 570 · see citing works
Oxytocin Attenuates Amygdala Reactivity to Fear in Generalized Social Anxiety Disorder
Labuschagne, I., Phan, K. L., Wood, A., Angstadt, M., Chua, P., Heinrichs, M., Stout, J. C., & Nathan, P. J. (2010) · Neuropsychopharmacology · doi:10.1038/npp.2010.123 · Cited by 549 · see citing works
Activity-dependent structural and functional plasticity of astrocyte-neuron interactions
Theodosis, D. T., Poulain, D. A., & Oliet, S. H. R. (2008) · Physiological Reviews · doi:10.1152/physrev.00036.2007 · Cited by 531 · see citing works
Effects of intranasal oxytocin on emotional face processing in women
Domes, G., Lischke, A., Berger, C., Grossmann, A., Hauenstein, K., Heinrichs, M., & Herpertz, S. C. (2010) · Psychoneuroendocrinology · doi:10.1016/j.psyneuen.2009.06.016 · Cited by 521 · see citing works
Effects of different kinds of couple interaction on cortisol and heart rate responses to stress in women
Ditzen, B., Neumann, I. D., Bodenmann, G., von Dawans, B., Turner, R. A., Ehlert, U., & Heinrichs, M. (2007) · Psychoneuroendocrinology · doi:10.1016/j.psyneuen.2007.03.011 · Cited by 525 · see citing works
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Increased brain and plasma oxytocin after nasal and peripheral administration in rats and mice
Neumann, I. D., Maloumby, R., Beiderbeck, D. I., Lukas, M., & Landgraf, R. (2013) · Psychoneuroendocrinology · doi:10.1016/j.psyneuen.2013.03.003 · Cited by 511 · see citing works
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Neuromodulation by Oxytocin and Vasopressin
Stoop, R. (2012) · Neuron · doi:10.1016/j.neuron.2012.09.025 · Cited by 506 · see citing works
Oxytocin in general anxiety and social fear: a translational approach
Neumann, I. D., & Slattery, D. A. (2016) · Biological Psychiatry · doi:10.1016/j.biopsych.2015.06.004 · Cited by 468 · see citing works
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Neuropeptides and social behaviour: effects of oxytocin and vasopressin in humans
HEINRICHS, M., & DOMES, G. (2008) · Progress in brain research · doi:10.1016/s0079-6123(08)00428-7 · Cited by 479 · see citing works
The hypothalamic-neurohypophysial system regulates the hypothalamic–pituitary–adrenal axis under stress: an old concept revisited
Engelmann, M., Landgraf, R., & Wotjak, C. T. (2004) · Frontiers in Neuroendocrinology · doi:10.1016/j.yfrne.2004.09.001 · Cited by 450 · see citing works
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Brain Oxytocin Correlates with Maternal Aggression: link to Anxiety
Bosch, O. J., Meddle, S. L., Beiderbeck, D. I., Douglas, A. J., & Neumann, I. D. (2005) · The Journal of Neuroscience · doi:10.1523/jneurosci.1342-05.2005 · Cited by 439 · see citing works
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Sex differences in oxytocin and vasopressin: Implications for autism spectrum disorders?
CARTER, C. (2007) · Behavioural Brain Research · doi:10.1016/j.bbr.2006.08.025 · Cited by 432 · see citing works
Acute effects of steroid hormones and neuropeptides on human social-emotional behavior: A review of single administration studies
Bos, P. A., Panksepp, J., Bluthé, R.-M., & Honk, J. V. (2012) · Frontiers in Neuroendocrinology · doi:10.1016/j.yfrne.2011.01.002 · Cited by 429 · see citing works
The Hypothalamic-Pituitary-Adrenal Axis: Development, Programming Actions of Hormones, and Maternal-Fetal Interactions
Sheng, J. A., Bales, N. J., Myers, S. A., Bautista, A. I., Roueinfar, M., Hale, T. M., & Handa, R. J. (2021) · Frontiers in Behavioral Neuroscience · doi:10.3389/fnbeh.2020.601939 · Cited by 418 · see citing works
Gonadal steroid hormones and the hypothalamo–pituitary–adrenal axis
Handa, R. J., & Weiser, M. J. (2014) · Frontiers in Neuroendocrinology · doi:10.1016/j.yfrne.2013.11.001 · Cited by 408 · see citing works
Oxytocin Makes a Face in Memory Familiar
Rimmele, U., Hediger, K., Heinrichs, M., & Klaver, P. (2009) · The Journal of Neuroscience · doi:10.1523/jneurosci.4260-08.2009 · Cited by 402 · see citing works
Both oxytocin and vasopressin are mediators of maternal care and aggression in rodents: From central release to sites of action
Bosch, O. J., & Neumann, I. D. (2012) · Hormones and Behavior · doi:10.1016/j.yhbeh.2011.11.002 · Cited by 400 · see citing works
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The neuropeptide oxytocin facilitates pro-social behavior and prevents social avoidance in rats and mice
Lukas, M., Toth, I., Reber, S. O., Slattery, D. A., Veenema, A. H., & Neumann, I. D. (2011) · Neuropsychopharmacology · doi:10.1038/npp.2011.95 · Cited by 403 · see citing works
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Is oxytocin “nature’s medicine”
Carter, C. S., Kenkel, W. M., MacLean, E. L., Wilson, S. R., Perkeybile, A. M., Yee, J. R., Ferris, C. F., Nazarloo, H. P., Porges, S. W., Davis, J. M., et al. (2020) · Pharmacological Reviews · doi:10.1124/pr.120.019398 · Cited by 394 · see citing works
Sniffing around oxytocin: Review and meta-analyses of trials in healthy and clinical groups with implications for pharmacotherapy
Bakermans-Kranenburg, M. J., & van IJzendoorn, M. H. (2013) · Translational Psychiatry · doi:10.1038/tp.2013.34 · Cited by 383 · see citing works
Oxytocin, vasopressin and sociality
CARTER, C., GRIPPO, A., POURNAJAFINAZARLOO, H., RUSCIO, M., & PORGES, S. (2008) · Progress in Brain Research · doi:10.1016/s0079-6123(08)00427-5 · Cited by 380 · see citing works
Intranasal oxytocin treatment for social deficits and biomarkers of response in children with autism
Parker, K. J., Oztan, O., Libove, R. A., Sumiyoshi, R. D., Jackson, L. P., Karhson, D. S., Summers, J. E., Hinman, K. E., Motonaga, K. S., Phillips, J. M., et al. (2017) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1705521114 · Cited by 349 · see citing works
The oxytocin receptor gene (OXTR) is associated with autism spectrum disorder: a meta-analysis
LoParo, D., & Waldman, I. D. (2015) · Molecular Psychiatry · doi:10.1038/mp.2014.77 · Cited by 343 · see citing works
Association between the oxytocin receptor (OXTR) gene and autism: relationship to Vineland Adaptive Behavior Scales and cognition
Lerer, E., Levi, S., Salomon, S., Darvasi, A., Yirmiya, N., & Ebstein, R. P. (2008) · Molecular Psychiatry · doi:10.1038/sj.mp.4002087 · Cited by 349 · see citing works
A review of safety, side-effects and subjective reactions to intranasal oxytocin in human research
MacDonald, E., Dadds, M. R., Brennan, J. L., Williams, K., Levy, F., & Cauchi, A. J. (2011) · Psychoneuroendocrinology · doi:10.1016/j.psyneuen.2011.02.015 · Cited by 352 · see citing works
Plasma oxytocin concentration during pregnancy is associated with development of postpartum depression
Skrundz, M., Bolten, M., Nast, I., Hellhammer, D. H., & Meinlschmidt, G. (2011) · Neuropsychopharmacology · doi:10.1038/npp.2011.74 · Cited by 349 · see citing works
Dopaminergic-neuropeptide interactions in the social brain
Skuse, D. H., & Gallagher, L. (2009) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2008.09.007 · Cited by 340 · see citing works
Central vasopressin and oxytocin release: regulation of complex social behaviours
VEENEMA, A., & NEUMANN, I. (2008) · Progress in Brain Research · doi:10.1016/s0079-6123(08)00422-6 · Cited by 338 · see citing works
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Oxytocin: Behavioral Associations and Potential as a Salivary Biomarker
SUE CARTER, C., POURNAJAFI‐NAZARLOO, H., KRAMER, K. M., ZIEGLER, T. E., WHITE‐TRAUT, R., BELLO, D., & SCHWERTZ, D. (2007) · Annals of the New York Academy of Sciences · doi:10.1196/annals.1384.006 · Cited by 323 · see citing works
Assembling the puzzle: pathways of oxytocin signaling in the brain
Grinevich, V., Knobloch-Bollmann, H. S., Eliava, M., Busnelli, M., & Chini, B. (2016) · Biological Psychiatry · doi:10.1016/j.biopsych.2015.04.013 · Cited by 309 · see citing works
Oxytocin and social perception: oxytocin increases perceived facial trustworthiness and attractiveness
Theodoridou, A., Rowe, A. C., Penton-Voak, I. S., & Rogers, P. J. (2009) · Hormones and Behavior · doi:10.1016/j.yhbeh.2009.03.019 · Cited by 318 · see citing works
Characterization of the oxytocin system regulating affiliative behavior in female prairie voles
Ross, H. E., Cole, C. D., Smith, Y., Neumann, I. D., Landgraf, R., Murphy, A. Z., & Young, L. J. (2009) · Neuroscience · doi:10.1016/j.neuroscience.2009.05.055 · Cited by 309 · see citing works
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Effects of intranasal oxytocin and vasopressin on cooperative behavior and associated brain activity in men
Rilling, J. K., DeMarco, A. C., Hackett, P. D., Thompson, R., Ditzen, B., Patel, R., & Pagnoni, G. (2012) · Psychoneuroendocrinology · doi:10.1016/j.psyneuen.2011.07.013 · Cited by 303 · see citing works
Social vocalizations can release oxytocin in humans
Seltzer, L. J., Ziegler, T. E., & Pollak, S. D. (2010) · Proceedings of the Royal Society B: Biological Sciences · doi:10.1098/rspb.2010.0567 · Cited by 298 · see citing works
Adaptive Responses of the Maternal Hypothalamic‐Pituitary‐Adrenal Axis during Pregnancy and Lactation
Brunton, P. J., Russell, J. A., & Douglas, A. J. (2008) · Journal of Neuroendocrinology · doi:10.1111/j.1365-2826.2008.01735.x · Cited by 298 · see citing works
Biased Oxytocinergic Modulation of Midbrain Dopamine Systems
Xiao, L., Priest, M. F., Nasenbeny, J., Lu, T., & Kozorovitskiy, Y. (2017) · Neuron · doi:10.1016/j.neuron.2017.06.003 · Cited by 292 · see citing works
Oxytocin, brain physiology, and functional connectivity: A review of intranasal oxytocin fMRI studies
Bethlehem, R. A. I., van Honk, J., Auyeung, B., & Baron-Cohen, S. (2013) · Psychoneuroendocrinology · doi:10.1016/j.psyneuen.2012.10.011 · Cited by 285 · see citing works
The regulation of social recognition, social communication and aggression: vasopressin in the social behavior neural network
Albers, H. E. (2012) · Hormones and Behavior · doi:10.1016/j.yhbeh.2011.10.007 · Cited by 288 · see citing works
Neuroanatomical evidence for reciprocal regulation of the corticotrophin-releasing factor and oxytocin systems in the hypothalamus and the bed nucleus of the stria terminalis of the rat: Implications for balancing stress and affect
Dabrowska, J., Hazra, R., Ahern, T. H., Guo, J.-D., McDonald, A. J., Mascagni, F., Muller, J. F., Young, L. J., & Rainnie, D. G. (2011) · Psychoneuroendocrinology · doi:10.1016/j.psyneuen.2011.03.003 · Cited by 274 · see citing works
The oxytocin receptor (OXTR) contributes to prosocial fund allocations in the dictator game and the social value orientations task
Israel, S., Lerer, E., Shalev, I., Uzefovsky, F., Riebold, M., Laiba, E., Bachner-Melman, R., Maril, A., Bornstein, G., Knafo, A., et al. (2009) · PLoS ONE · doi:10.1371/journal.pone.0005535 · Cited by 278 · see citing works
Evolution of oxytocin pathways in the brain of vertebrates
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The animal and human neuroendocrinology of social cognition, motivation and behavior
McCall, C., & Singer, T. (2012) · Nature Neuroscience · doi:10.1038/nn.3084 · Cited by 273 · see citing works
Opposite effects of maternal separation on intermale and maternal aggression in C57BL/6 mice: Link to hypothalamic vasopressin and oxytocin immunoreactivity
Veenema, A. H., Bredewold, R., & Neumann, I. D. (2007) · Psychoneuroendocrinology · doi:10.1016/j.psyneuen.2007.02.008 · Cited by 263 · see citing works
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Post-learning intranasal oxytocin modulates human memory for facial identity
Savaskan, E., Ehrhardt, R., Schulz, A., Walter, M., & Schächinger, H. (2008) · Psychoneuroendocrinology · doi:10.1016/j.psyneuen.2007.12.004 · Cited by 262 · see citing works
Centrally released oxytocin mediates mating-induced anxiolysis in male rats
Waldherr, M., & Neumann, I. D. (2007) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0705860104 · Cited by 261 · see citing works
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Authors: Landgraf, R., & Neumann, I. D.
Affiliations: Max Planck Society (DE)
University of Regensburg (DE)
Corresponding author: Landgraf, Rainer
ORCID: Neumann, Inga D.
Venue: Frontiers in Neuroendocrinology
Publisher: Elsevier BV
Published: 2004-09
Volume: 25
Issue: 3-4
Pages: 150-176
ISSN: 0091-3022 (ISSN-L)
DOI: 10.1016/j.yfrne.2004.05.001
OpenAlex: W1996368170
Cited by: 1,059
References: 295
Citations by year: 2026: 24 · 2025: 25 · 2024: 38 · 2023: 38 · 2022: 48 · 2021: 52 · 2020: 61 · 2019: 47 · 2018: 54 · 2017: 59 · 2016: 60 · 2015: 54 · 2014: 61 · 2013: 66 · 2012: 61
FWCI: 43.49
Type: journal article
Language: en
Open access: closed
Licence: www.elsevier.com/tdm/userlicense/1.0/ (tdm)
Locations: Frontiers in Neuroendocrinology (publishedVersion)
PubMed (publishedVersion)
Max Planck Institute for Plasma Physics (submittedVersion)
University of Regensburg Publication Server (University of Regensburg) (submittedVersion)
MPG.PuRe (Max Planck Society) (submittedVersion)
Topics: Social Sciences › Psychology › Social Psychology › Neuroendocrine regulation and behavior
Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuropeptides and Animal Physiology
Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
Keywords: vasopressin, oxytocin, neuropeptide release, dendritic release, neurosecretory cells, intracellular calcium stores, synaptic input, transcription factors, neuromodulation, neuroendocrine regulation, psychopathology
MeSH terms: Animals, Brain, Cell Communication, Humans, Nerve Net, Neurons, Oxytocin, Vasopressins, Feedback, Physiological

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Vasopressin and oxytocin release within the brain: a dynamic concept of multiple and variable modes of neuropeptide communication

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