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As discussed in this Review, a cilium is an elongated and highly specialized structure, and, together with its ability to selectively traffic and concentrate proteins, lipids and second messengers, it creates a signaling environment distinct from the cell body. Ciliary signaling pathways adopt a bow-tie network architecture, in which diverse inputs converge on shared effectors and second messengers before diverging to multiple outputs. Unlike other cellular bow-tie systems, cells exploit ciliary geometry, compartmentalization and infrastructure to enhance sensitivity at multiple scales, from individual molecular reactions to entire signaling pathways. In cilia, integration of the bow-tie network architecture with their specialized structure and unique environment confers robustness and evolvability, which enables cilia to acquire diverse signaling roles. However, this versatility comes with vulnerability \u2013 rare mutations that disrupt the features most essential for cilia robustness cause multisystem ciliopathies.<\/jats:p>","DOI":"10.1242\/jcs.264325","type":"journal-article","created":{"date-parts":[[2025,10,24]],"date-time":"2025-10-24T12:36:57Z","timestamp":1761309417000},"source":"Crossref","is-referenced-by-count":15,"title":["Design principles of ciliary signaling"],"prefix":"10.1242","volume":"138","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5246-1229","authenticated-orcid":false,"given":"Carolyn M.","family":"Ott","sequence":"first","affiliation":[{"name":"Janelia Research Campus, Howard Hughes Medical Institute , Ashburn, VA 20147 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8601-3501","authenticated-orcid":false,"given":"Jennifer","family":"Lippincott-Schwartz","sequence":"additional","affiliation":[{"name":"Janelia Research Campus, Howard Hughes Medical Institute , Ashburn, VA 20147 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2025,10,24]]},"reference":[{"key":"2025102408365148800_JCS264325C1","first-page":"198","article-title":"Reduction of dimensionality in biological diffusion processes","volume-title":"Structural Chemistry and Molecular Biology","author":"Adam","year":"1968"},{"key":"2025102408365148800_JCS264325C2","doi-asserted-by":"publisher","first-page":"ar138","DOI":"10.1091\/mbc.E24-05-0226","article-title":"Spatial and signaling overlap of growth factor receptor systems at clathrin-coated sites","volume":"35","author":"Alfonzo-M\u00e9ndez","year":"2024","journal-title":"Mol. 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