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The parent neurons of the axons are in the K\u00f6lliker\u2013Fuse nucleus, and their terminals form calyx-like multirelease-site synapses in the rodent forebrain extended amygdala similar to the calyx of Held in the auditory brain stem. The latter is a giant, excitatory, cup-like axo-somatic high-fidelity synapse. The PACAP-positive terminals also form enveloping axo-somatic specialization with mixed glutamatergic and cholinergic molecular identities, co-expressing vesicular glutamate transporter 1 (VGluT1), VGluT2, vesicular acetylcholine transport (VAChT), and the neuropeptides PACAP, calcitonin gene\u2013related peptide, and neurotensin, together with calretinin in the presynaptic compartment. We identified a distinct neuronal subpopulation in the pontine K\u00f6lliker\u2013Fuse region of the parabrachial complex that gives rise to these calyceal terminals, which engulf Protein kinase C delta (PKC\u03b4+) \/ Glutamate delta receptor 1 (GluD1+) somata in the capsular central amygdala and oval bed nucleus of the stria terminalis. Strikingly, GluD1 immunolabeling is concentrated at axo-somatic contact zones apposed to VAChT+ presynaptic vesicle cluster zones but is absent from postsynaptic densities of conventional type I synapses within the same terminals. The results demonstrate a previously unrecognized multimodal calyx-like synapse in the forebrain, with parallel fast ionotropic and modulatory peptidergic neurotransmission mechanisms, which is a substrate for high-fidelity signal transmission within viscerosensory-emotional circuits.<\/jats:p>","DOI":"10.1093\/pnasnexus\/pgag083","type":"journal-article","created":{"date-parts":[[2026,3,23]],"date-time":"2026-03-23T07:57:12Z","timestamp":1774252632000},"source":"Crossref","is-referenced-by-count":2,"title":["A multitransmitter, synaptic specification\u2013segregated calyx-like synapse linking pontine pituitary adenylate cyclase\u2013activating polypeptide neurons to forebrain-extended amygdala"],"prefix":"10.1093","volume":"5","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7422-5136","authenticated-orcid":false,"given":"Limei","family":"Zhang","sequence":"first","affiliation":[{"name":"Department of Physiology, School of Medicine, National Autonomous University of Mexico (UNAM), 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Microscopy Core, National Institute on Drug Abuse, NIH , Baltimore, MD 21224 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shiliang","family":"Zhang","sequence":"additional","affiliation":[{"name":"Confocal and Electron Microscopy Core, National Institute on Drug Abuse, NIH , Baltimore, MD 21224 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-3554-2356","authenticated-orcid":false,"given":"Martin K H","family":"Sch\u00e4fer","sequence":"additional","affiliation":[{"name":"Center for Mind, Brain and Behaviour and Department of Medical Cell Biology, Institute for Anatomy and Cell Biology, Philipps University of Marburg , Marburg 35037 ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0987-0785","authenticated-orcid":false,"given":"Rafael A","family":"Barrio","sequence":"additional","affiliation":[{"name":"Department of 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