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Decades of elegant research have illuminated the molecular mechanisms and machinery that support the release of neurotransmitters via synaptic vesicle exocytosis, as well as the secretion of signal\u2010peptide bearing proteins through the endoplasmic reticulum (ER)\u2010Golgi based secretory pathways. However, it is now increasingly appreciated that signal\u2010peptide lacking \u201cleaderless\u201d proteins can also be secreted via ER\u2010Golgi\u2010independent mechanisms collectively termed unconventional protein secretion (UcPS). In this review, we highlight the physiological and pathological consequences of UcPS in the central nervous system. UcPS supports the secretion of aggregation\u2010prone proteins such as \u03b1\u2010synuclein and mutant huntingtin, pro\u2010inflammatory mediators including interleukin\u20101\u03b2 and high mobility group box protein 1, and neuroprotective or angiogenic factors such as fibroblast growth factor 2. Furthermore, several retroelement\u2010derived proteins, encoded by ancient genomic elements with structural homology to retroviruses, are also secreted via unconventional pathways, and are thought to regulate essential CNS processes such as synaptic plasticity. These diverse cargoes underscore the functional range of UcPS in neuronal and glial biology. We summarize current understanding of the major UcPS pathways used by CNS cells. These mechanisms include plasma\u2010membrane pore\u2010mediated release facilitated by proteins such as gasdermin\u2010D, as well as vesicular routes in which UcPS cargoes enter organelles of the autophagic and endolysosomal systems that subsequently fuse with the plasma membrane to enable extracellular release. Finally, we discuss key unresolved questions regarding the regulation of UcPS, including the molecular features that target select leaderless cargoes toward UcPS, how the balance between conventional secretion and UcPS shifts under cellular stress, and the current understanding of the diverse molecular machinery that mediates the vesicular form of UcPS.<\/jats:p>\n                  <jats:p>\n                    <jats:boxed-text content-type=\"graphic\" position=\"anchor\">\n                      <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" mimetype=\"image\/png\" position=\"anchor\" specific-use=\"enlarged-web-image\" xlink:href=\"graphic\/jnc70532-gra-0001-m.png\">\n                        <jats:alt-text>image<\/jats:alt-text>\n                      <\/jats:graphic>\n                    <\/jats:boxed-text>\n                  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