{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T08:55:10Z","timestamp":1790585710355,"version":"4.1.0"},"reference-count":107,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"funder":[{"DOI":"10.13039\/100000154","name":"National Science Foundation Division of Integrative Organismal Systems","doi-asserted-by":"publisher","award":["1625154"],"award-info":[{"award-number":["1625154"]}],"id":[{"id":"10.13039\/100000154","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Developmental Biology"],"published-print":{"date-parts":[[2026,11]]},"DOI":"10.1016\/j.ydbio.2026.08.012","type":"journal-article","created":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T14:46:31Z","timestamp":1787669191000},"page":"141-148","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"special_numbering":"C","title":["Shedding light on the proteolytically released ectodomains of membrane-bound Semaphorins"],"prefix":"10.1016","volume":"539","author":[{"given":"Collin M.","family":"MacLeod","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Riley M.","family":"St Clair","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7650-3817","authenticated-orcid":false,"given":"Bryan A.","family":"Ballif","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alicia M.","family":"Ebert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"issue":"819","key":"10.1016\/j.ydbio.2026.08.012_bib1","doi-asserted-by":"crossref","first-page":"eadh7673","DOI":"10.1126\/scisignal.adh7673","article-title":"Balancing act of small GTPases downstream of plexin-A4 signaling motifs promotes dendrite elaboration in mammalian cortical neurons","volume":"17","author":"Abushalbaq","year":"2024","journal-title":"Sci. Signal."},{"key":"10.1016\/j.ydbio.2026.08.012_bib2","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/978-1-4939-6448-2_1","article-title":"Semaphorins and their signaling mechanisms","volume":"1493","author":"Alto","year":"2017","journal-title":"Methods Mol. Biol."},{"key":"10.1016\/j.ydbio.2026.08.012_bib3","doi-asserted-by":"crossref","DOI":"10.1016\/j.molmet.2023.101731","article-title":"Semaphorin 4B is an ADAM17-cleaved adipokine that inhibits adipocyte differentiation and thermogenesis","volume":"73","author":"Amin","year":"2023","journal-title":"Mol. Metabol."},{"issue":"19","key":"10.1016\/j.ydbio.2026.08.012_bib4","doi-asserted-by":"crossref","first-page":"3709","DOI":"10.1242\/dev.112185","article-title":"Semaphorin 6B acts as a receptor in post-crossing commissural axon guidance","volume":"141","author":"Andermatt","year":"2014","journal-title":"Development"},{"issue":"9","key":"10.1016\/j.ydbio.2026.08.012_bib5","doi-asserted-by":"crossref","DOI":"10.3390\/ijms20092144","article-title":"Multifaceted functional role of semaphorins in glioblastoma","volume":"20","author":"Angelucci","year":"2019","journal-title":"Int. J. Mol. Sci."},{"key":"10.1016\/j.ydbio.2026.08.012_bib6","doi-asserted-by":"crossref","first-page":"19","DOI":"10.3389\/fnins.2015.00019","article-title":"Function and regulation of Rnd proteins in cortical projection neuron migration","volume":"9","author":"Azzarelli","year":"2015","journal-title":"Front. Neurosci."},{"issue":"15","key":"10.1016\/j.ydbio.2026.08.012_bib7","doi-asserted-by":"crossref","first-page":"5212","DOI":"10.1158\/0008-5472.CAN-04-0126","article-title":"Class IV semaphorins promote angiogenesis by stimulating Rho-initiated pathways through plexin-B","volume":"64","author":"Basile","year":"2004","journal-title":"Cancer Res."},{"issue":"24","key":"10.1016\/j.ydbio.2026.08.012_bib8","doi-asserted-by":"crossref","first-page":"9017","DOI":"10.1073\/pnas.0508825103","article-title":"Semaphorin 4D provides a link between axon guidance processes and tumor-induced angiogenesis","volume":"103","author":"Basile","year":"2006","journal-title":"Proc. Natl. Acad. Sci. U. S. A"},{"issue":"9","key":"10.1016\/j.ydbio.2026.08.012_bib9","doi-asserted-by":"crossref","first-page":"6899","DOI":"10.1074\/jbc.M609570200","article-title":"MT1-MMP controls tumor-induced angiogenesis through the release of semaphorin 4D","volume":"282","author":"Basile","year":"2007","journal-title":"J. Biol. Chem."},{"issue":"8","key":"10.1016\/j.ydbio.2026.08.012_bib10","doi-asserted-by":"crossref","first-page":"1609","DOI":"10.1007\/s00018-016-2137-x","article-title":"Transmembrane semaphorins, forward and reverse signaling: have a look both ways","volume":"73","author":"Battistini","year":"2016","journal-title":"Cell. Mol. Life Sci."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib11","doi-asserted-by":"crossref","first-page":"7059","DOI":"10.1038\/s41467-024-51009-1","article-title":"Semaphorin heterodimerization in cis regulates membrane targeting and neocortical wiring","volume":"15","author":"Bessa","year":"2024","journal-title":"Nat. Commun."},{"issue":"2","key":"10.1016\/j.ydbio.2026.08.012_bib12","doi-asserted-by":"crossref","first-page":"318","DOI":"10.4049\/jimmunol.148.2.318","article-title":"Increased surface expression of a newly identified 150-kDa dimer early after human T lymphocyte activation","volume":"148","author":"Bougeret","year":"1992","journal-title":"J. Immunol."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib13","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1111\/j.1471-4159.2012.07885.x","article-title":"Transmembrane semaphorin5B is proteolytically processed into a repulsive neural guidance cue","volume":"123","author":"Browne","year":"2012","journal-title":"J. Neurochem."},{"issue":"17","key":"10.1016\/j.ydbio.2026.08.012_bib14","doi-asserted-by":"crossref","first-page":"3821","DOI":"10.1016\/j.febslet.2005.05.079","article-title":"Semaphorin 4B interacts with the post-synaptic density protein PSD-95\/SAP90 and is recruited to synapses through a C-terminal PDZ-binding motif","volume":"579","author":"Burkhardt","year":"2005","journal-title":"FEBS Lett."},{"issue":"22","key":"10.1016\/j.ydbio.2026.08.012_bib15","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2400648121","article-title":"Astrocyte-to-microglia communication via Sema4B-Plexin-B2 modulates injury-induced reactivity of microglia","volume":"121","author":"Casden","year":"2024","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"issue":"3","key":"10.1016\/j.ydbio.2026.08.012_bib16","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1016\/j.celrep.2017.09.074","article-title":"Phosphorylation of iRhom2 controls stimulated proteolytic shedding by the metalloprotease ADAM17\/TACE","volume":"21","author":"Cavadas","year":"2017","journal-title":"Cell Rep."},{"issue":"4","key":"10.1016\/j.ydbio.2026.08.012_bib17","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1093\/intimm\/dxh224","article-title":"Soluble CD100 functions on human monocytes and immature dendritic cells require plexin C1 and plexin B1, respectively","volume":"17","author":"Chabbert-de Ponnat","year":"2005","journal-title":"Int. Immunol."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib18","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1038\/nrm1788","article-title":"Function and regulation of Rnd proteins","volume":"7","author":"Chardin","year":"2006","journal-title":"Nat. Rev. Mol. Cell Biol."},{"issue":"Suppl. 4","key":"10.1016\/j.ydbio.2026.08.012_bib19","doi-asserted-by":"crossref","first-page":"S468","DOI":"10.21037\/tlcr.2019.12.22","article-title":"Importance of semaphorins in cancer immunity","volume":"8","author":"Chen","year":"2019","journal-title":"Transl. Lung Cancer Res."},{"issue":"11","key":"10.1016\/j.ydbio.2026.08.012_bib20","doi-asserted-by":"crossref","first-page":"4321","DOI":"10.1182\/blood-2004-07-2885","article-title":"Sema4D induces angiogenesis through Met recruitment by Plexin B1","volume":"105","author":"Conrotto","year":"2005","journal-title":"Blood"},{"issue":"7","key":"10.1016\/j.ydbio.2026.08.012_bib21","doi-asserted-by":"crossref","first-page":"4348","DOI":"10.4049\/jimmunol.166.7.4348","article-title":"Biological activity of soluble CD100. II. Soluble CD100, similarly to H-SemaIII, inhibits immune cell migration","volume":"166","author":"Delaire","year":"2001","journal-title":"J. Immunol."},{"issue":"23","key":"10.1016\/j.ydbio.2026.08.012_bib22","doi-asserted-by":"crossref","first-page":"6333","DOI":"10.1523\/JNEUROSCI.5381-06.2007","article-title":"Plexin-B2, but not Plexin-B1, critically modulates neuronal migration and patterning of the developing nervous system in vivo","volume":"27","author":"Deng","year":"2007","journal-title":"J. Neurosci."},{"issue":"6","key":"10.1016\/j.ydbio.2026.08.012_bib23","doi-asserted-by":"crossref","first-page":"659","DOI":"10.4161\/cbt.4.6.1733","article-title":"Recombinant semaphorin 6A-1 ectodomain inhibits in vivo growth factor and tumor cell line-induced angiogenesis","volume":"4","author":"Dhanabal","year":"2005","journal-title":"Cancer Biol. Ther."},{"issue":"14","key":"10.1016\/j.ydbio.2026.08.012_bib24","doi-asserted-by":"crossref","DOI":"10.1242\/jcs.261469","article-title":"The intracellular domain of Sema6A is essential for development of the zebrafish retina","volume":"137","author":"Dumas","year":"2024","journal-title":"J. Cell Sci."},{"issue":"12","key":"10.1016\/j.ydbio.2026.08.012_bib25","doi-asserted-by":"crossref","first-page":"2473","DOI":"10.1242\/dev.103499","article-title":"Sema6a and Plxna2 mediate spatially regulated repulsion within the developing eye to promote eye vesicle cohesion","volume":"141","author":"Ebert","year":"2014","journal-title":"Development"},{"issue":"7","key":"10.1016\/j.ydbio.2026.08.012_bib26","doi-asserted-by":"crossref","first-page":"4341","DOI":"10.4049\/jimmunol.166.7.4341","article-title":"Biological activity of soluble CD100. I. The extracellular region of CD100 is released from the surface of T lymphocytes by regulated proteolysis","volume":"166","author":"Elhabazi","year":"2001","journal-title":"J. Immunol."},{"issue":"7","key":"10.1016\/j.ydbio.2026.08.012_bib27","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1002\/dvdy.24512","article-title":"Identification of target genes downstream of semaphorin6A\/PlexinA2 signaling in zebrafish","volume":"246","author":"Emerson","year":"2017","journal-title":"Dev. Dyn."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib28","doi-asserted-by":"crossref","first-page":"e15","DOI":"10.1182\/blood-2010-05-283838","article-title":"Deciphering the human platelet sheddome","volume":"117","author":"Fong","year":"2011","journal-title":"Blood"},{"issue":"9","key":"10.1016\/j.ydbio.2026.08.012_bib29","doi-asserted-by":"crossref","first-page":"865","DOI":"10.1038\/embor.2008.139","article-title":"Tyrosine phosphorylation in semaphorin signalling: shifting into overdrive","volume":"9","author":"Franco","year":"2008","journal-title":"EMBO Rep."},{"issue":"9","key":"10.1016\/j.ydbio.2026.08.012_bib30","doi-asserted-by":"crossref","first-page":"720","DOI":"10.1038\/ncb843","article-title":"The semaphorin 4D receptor controls invasive growth by coupling with Met","volume":"4","author":"Giordano","year":"2002","journal-title":"Nat. Cell Biol."},{"issue":"5","key":"10.1016\/j.ydbio.2026.08.012_bib31","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1016\/S0092-8674(00)80766-7","article-title":"Unified nomenclature for the semaphorins\/collapsins. Semaphorin Nomenclature committee","volume":"97","author":"Goodman","year":"1999","journal-title":"Cell"},{"issue":"6","key":"10.1016\/j.ydbio.2026.08.012_bib32","doi-asserted-by":"crossref","first-page":"1461","DOI":"10.1002\/art.38425","article-title":"Secreted semaphorin 5A activates immune effector cells and is a biomarker for rheumatoid arthritis","volume":"66","author":"Gras","year":"2014","journal-title":"Arthritis Rheumatol."},{"key":"10.1016\/j.ydbio.2026.08.012_bib33","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.23968","article-title":"Phosphorylation of iRhom2 at the plasma membrane controls mammalian TACE-dependent inflammatory and growth factor signalling","volume":"6","author":"Grieve","year":"2017","journal-title":"eLife"},{"issue":"19","key":"10.1016\/j.ydbio.2026.08.012_bib34","doi-asserted-by":"crossref","first-page":"3685","DOI":"10.1007\/s00018-014-1633-0","article-title":"Eph- and ephrin-dependent mechanisms in tumor and stem cell dynamics","volume":"71","author":"Gucciardo","year":"2014","journal-title":"Cell. Mol. Life Sci."},{"issue":"7","key":"10.1016\/j.ydbio.2026.08.012_bib36","doi-asserted-by":"crossref","first-page":"1259","DOI":"10.1038\/s41418-018-0097-4","article-title":"Sema4C\/PlexinB2 signaling controls breast cancer cell growth, hormonal dependence and tumorigenic potential","volume":"25","author":"Gurrapu","year":"2018","journal-title":"Cell Death Differ."},{"issue":"6","key":"10.1016\/j.ydbio.2026.08.012_bib35","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1080\/19336918.2016.1197479","article-title":"Transmembrane semaphorins: multimodal signaling cues in development and cancer","volume":"10","author":"Gurrapu","year":"2016","journal-title":"Cell Adhes. Migrat."},{"issue":"15","key":"10.1016\/j.ydbio.2026.08.012_bib37","doi-asserted-by":"crossref","first-page":"2635","DOI":"10.1038\/emboj.2010.147","article-title":"Cis interaction between Semaphorin6A and Plexin-A4 modulates the repulsive response to Sema6A","volume":"29","author":"Haklai-Topper","year":"2010","journal-title":"EMBO J."},{"key":"10.1016\/j.ydbio.2026.08.012_bib38","doi-asserted-by":"crossref","DOI":"10.3389\/fcell.2025.1557653","article-title":"Emerging roles for E3 ubiquitin ligases in neural development and disease","volume":"13","author":"Hale","year":"2025","journal-title":"Front. Cell Dev. Biol."},{"key":"10.1016\/j.ydbio.2026.08.012_bib39","doi-asserted-by":"crossref","first-page":"1744","DOI":"10.1002\/1873-3468.70293","article-title":"ABL drives phosphorylation of Semaphorin 6A at conserved tyrosines necessary for zebrafish retinal maintenance","volume":"600","author":"Haney","year":"2026","journal-title":"FEBS Lett."},{"key":"10.1016\/j.ydbio.2026.08.012_bib40","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1016\/j.isci.2019.10.034","article-title":"Semaphorin 6A-Plexin A2\/A4 interactions with radial glia regulate migration termination of superficial layer cortical neurons","volume":"21","author":"Hatanaka","year":"2019","journal-title":"iScience"},{"issue":"12","key":"10.1016\/j.ydbio.2026.08.012_bib41","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0008477","article-title":"Identification of beta-secretase (BACE1) substrates using quantitative proteomics","volume":"4","author":"Hemming","year":"2009","journal-title":"PLoS One"},{"issue":"12","key":"10.1016\/j.ydbio.2026.08.012_bib42","doi-asserted-by":"crossref","first-page":"5262","DOI":"10.4049\/jimmunol.157.12.5262","article-title":"CD100 is associated with CD45 at the surface of human T lymphocytes. Role in T cell homotypic adhesion","volume":"157","author":"Herold","year":"1996","journal-title":"J. Immunol."},{"issue":"3","key":"10.1016\/j.ydbio.2026.08.012_bib43","doi-asserted-by":"crossref","first-page":"270","DOI":"10.1046\/j.1365-3083.2002.01129.x","article-title":"Sema7A is a potent monocyte stimulator","volume":"56","author":"Holmes","year":"2002","journal-title":"Scand. J. Immunol."},{"issue":"22","key":"10.1016\/j.ydbio.2026.08.012_bib44","doi-asserted-by":"crossref","first-page":"3765","DOI":"10.1007\/s00018-012-1019-0","article-title":"Plexin structures are coming: opportunities for multilevel investigations of semaphorin guidance receptors, their cell signaling mechanisms, and functions","volume":"69","author":"Hota","year":"2012","journal-title":"Cell. Mol. Life Sci."},{"issue":"8","key":"10.1016\/j.ydbio.2026.08.012_bib45","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1002\/cm.20527","article-title":"Extracellular inhibitors, repellents, and semaphorin\/plexin\/MICAL-mediated actin filament disassembly","volume":"68","author":"Hung","year":"2011","journal-title":"Cytoskeleton (Hoboken)"},{"issue":"12","key":"10.1016\/j.ydbio.2026.08.012_bib46","doi-asserted-by":"crossref","first-page":"9174","DOI":"10.1074\/jbc.M009051200","article-title":"Sema4c, a transmembrane semaphorin, interacts with a post-synaptic density protein, PSD-95","volume":"276","author":"Inagaki","year":"2001","journal-title":"J. Biol. Chem."},{"issue":"12","key":"10.1016\/j.ydbio.2026.08.012_bib47","doi-asserted-by":"crossref","first-page":"1750","DOI":"10.1111\/jcmm.17416","article-title":"Soluble Sema4D cleaved from osteoclast precursors by TACE suppresses osteoblastogenesis","volume":"27","author":"Ishii","year":"2023","journal-title":"J. Cell Mol. Med."},{"issue":"35","key":"10.1016\/j.ydbio.2026.08.012_bib48","doi-asserted-by":"crossref","first-page":"12343","DOI":"10.1074\/jbc.RA120.013758","article-title":"Negatively charged amino acids in the stalk region of membrane proteins reduce ectodomain shedding","volume":"295","author":"Iwagishi","year":"2020","journal-title":"J. Biol. Chem."},{"issue":"6","key":"10.1016\/j.ydbio.2026.08.012_bib49","doi-asserted-by":"crossref","first-page":"1100","DOI":"10.1111\/j.1538-7836.2012.04708.x","article-title":"Semaphorin 7A inhibits platelet production from CD34+ progenitor cells","volume":"10","author":"Jaimes","year":"2012","journal-title":"J. Thromb. Haemostasis : JTH"},{"issue":"7319","key":"10.1016\/j.ydbio.2026.08.012_bib50","doi-asserted-by":"crossref","first-page":"1118","DOI":"10.1038\/nature09468","article-title":"Structural basis of semaphorin-plexin signalling","volume":"467","author":"Janssen","year":"2010","journal-title":"Nature"},{"issue":"12","key":"10.1016\/j.ydbio.2026.08.012_bib51","doi-asserted-by":"crossref","first-page":"1293","DOI":"10.1038\/nsmb.2416","article-title":"Neuropilins lock secreted semaphorins onto plexins in a ternary signaling complex","volume":"19","author":"Janssen","year":"2012","journal-title":"Nat. Struct. Mol. Biol."},{"issue":"17","key":"10.1016\/j.ydbio.2026.08.012_bib52","doi-asserted-by":"crossref","first-page":"3292","DOI":"10.1242\/dev.105544","article-title":"Semaphorin signalling during development","volume":"141","author":"Jongbloets","year":"2014","journal-title":"Development"},{"issue":"50","key":"10.1016\/j.ydbio.2026.08.012_bib53","doi-asserted-by":"crossref","first-page":"39647","DOI":"10.1074\/jbc.M006316200","article-title":"The orthologous human and murine semaphorin 6A-1 proteins (SEMA6A-1\/Sema6A-1) bind to the enabled\/vasodilator-stimulated phosphoprotein-like protein (EVL) via a novel carboxyl-terminal zyxin-like domain","volume":"275","author":"Klostermann","year":"2000","journal-title":"J. Biol. Chem."},{"issue":"5","key":"10.1016\/j.ydbio.2026.08.012_bib54","doi-asserted-by":"crossref","first-page":"831","DOI":"10.1016\/0896-6273(92)90237-8","article-title":"Fasciclin IV: sequence, expression, and function during growth cone guidance in the grasshopper embryo","volume":"9","author":"Kolodkin","year":"1992","journal-title":"Neuron"},{"issue":"3","key":"10.1016\/j.ydbio.2026.08.012_bib55","doi-asserted-by":"crossref","first-page":"548","DOI":"10.1016\/j.neuron.2016.06.018","article-title":"Structural basis for plexin activation and regulation","volume":"91","author":"Kong","year":"2016","journal-title":"Neuron"},{"key":"10.1016\/j.ydbio.2026.08.012_bib56","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1016\/S0065-2776(03)81005-2","article-title":"Roles of the semaphorin family in immune regulation","volume":"81","author":"Kumanogoh","year":"2003","journal-title":"Adv. Immunol."},{"key":"10.1016\/j.ydbio.2026.08.012_bib57","doi-asserted-by":"crossref","DOI":"10.1016\/j.jchemneu.2023.102318","article-title":"Cleavage of semaphorin 4 C interferes with the neuroprotective effect of the semaphorin 4 C\/Plexin B2 pathway on experimental intracerebral hemorrhage in rats","volume":"132","author":"Li","year":"2023","journal-title":"J. Chem. Neuroanat."},{"issue":"35","key":"10.1016\/j.ydbio.2026.08.012_bib58","doi-asserted-by":"crossref","first-page":"12353","DOI":"10.1074\/jbc.H120.015304","article-title":"To cut or not to cut: new rules for proteolytic shedding of membrane proteins","volume":"295","author":"Lichtenthaler","year":"2020","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.ydbio.2026.08.012_bib59","series-title":"Greek-English Lexicon","author":"Lidell","year":"1940"},{"issue":"5","key":"10.1016\/j.ydbio.2026.08.012_bib60","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0064265","article-title":"Increased levels of plasma soluble Sema4D in patients with heart failure","volume":"8","author":"Lu","year":"2013","journal-title":"PLoS One"},{"issue":"28","key":"10.1016\/j.ydbio.2026.08.012_bib61","doi-asserted-by":"crossref","first-page":"11433","DOI":"10.1073\/pnas.1302553110","article-title":"iRhom2 controls the substrate selectivity of stimulated ADAM17-dependent ectodomain shedding","volume":"110","author":"Maretzky","year":"2013","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"issue":"9","key":"10.1016\/j.ydbio.2026.08.012_bib62","doi-asserted-by":"crossref","first-page":"821","DOI":"10.1111\/j.1365-2443.2004.00766.x","article-title":"Sema4D stimulates axonal outgrowth of embryonic DRG sensory neurones","volume":"9","author":"Masuda","year":"2004","journal-title":"Genes Cells"},{"issue":"4","key":"10.1016\/j.ydbio.2026.08.012_bib63","doi-asserted-by":"crossref","first-page":"655","DOI":"10.1016\/j.neuron.2012.12.031","article-title":"Interaxonal interaction defines tiled presynaptic innervation in C. elegans","volume":"77","author":"Mizumoto","year":"2013","journal-title":"Neuron"},{"issue":"20","key":"10.1016\/j.ydbio.2026.08.012_bib64","doi-asserted-by":"crossref","first-page":"4221","DOI":"10.1182\/blood-2012-11-470609","article-title":"Identification of a calmodulin-binding domain in Sema4D that regulates its exodomain shedding in platelets","volume":"121","author":"Mou","year":"2013","journal-title":"Blood"},{"issue":"7","key":"10.1016\/j.ydbio.2026.08.012_bib65","doi-asserted-by":"crossref","first-page":"3630","DOI":"10.1128\/JVI.03634-14","article-title":"Viral semaphorin inhibits dendritic cell phagocytosis and migration but is not essential for gammaherpesvirus-induced lymphoproliferation in malignant catarrhal fever","volume":"89","author":"Myster","year":"2015","journal-title":"J. Virol."},{"issue":"11","key":"10.1016\/j.ydbio.2026.08.012_bib66","doi-asserted-by":"crossref","first-page":"1473","DOI":"10.1038\/nm.2489","article-title":"Suppression of bone formation by osteoclastic expression of semaphorin 4D","volume":"17","author":"Negishi-Koga","year":"2011","journal-title":"Nat. Med."},{"issue":"7319","key":"10.1016\/j.ydbio.2026.08.012_bib67","doi-asserted-by":"crossref","first-page":"1123","DOI":"10.1038\/nature09473","article-title":"Structural basis for semaphorin signalling through the plexin receptor","volume":"467","author":"Nogi","year":"2010","journal-title":"Nature"},{"key":"10.1016\/j.ydbio.2026.08.012_bib68","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1186\/1749-8104-4-18","article-title":"Semaphorin 5B mediates synapse elimination in hippocampal neurons","volume":"4","author":"O'Connor","year":"2009","journal-title":"Neural Dev."},{"issue":"28","key":"10.1016\/j.ydbio.2026.08.012_bib69","doi-asserted-by":"crossref","first-page":"25671","DOI":"10.1074\/jbc.M303047200","article-title":"Direct interaction of Rnd1 with Plexin-B1 regulates PDZ-RhoGEF-mediated Rho activation by Plexin-B1 and induces cell contraction in COS-7 cells","volume":"278","author":"Oinuma","year":"2003","journal-title":"J. Biol. Chem."},{"issue":"36","key":"10.1016\/j.ydbio.2026.08.012_bib70","doi-asserted-by":"crossref","first-page":"28200","DOI":"10.1074\/jbc.M110.147546","article-title":"Semaphorin 4D\/Plexin-B1 stimulates PTEN activity through R-Ras GTPase-activating protein activity, inducing growth cone collapse in hippocampal neurons","volume":"285","author":"Oinuma","year":"2010","journal-title":"J. Biol. Chem."},{"issue":"6947","key":"10.1016\/j.ydbio.2026.08.012_bib71","doi-asserted-by":"crossref","first-page":"398","DOI":"10.1038\/nature01790","article-title":"Semaphorin 7A promotes axon outgrowth through integrins and MAPKs","volume":"424","author":"Pasterkamp","year":"2003","journal-title":"Nature"},{"issue":"7","key":"10.1016\/j.ydbio.2026.08.012_bib72","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0158686","article-title":"Reverse signaling by Semaphorin-6A regulates cellular aggregation and neuronal morphology","volume":"11","author":"Perez-Branguli","year":"2016","journal-title":"PLoS One"},{"key":"10.1016\/j.ydbio.2026.08.012_bib73","doi-asserted-by":"crossref","DOI":"10.1016\/j.lfs.2024.123281","article-title":"Semaphorins and its receptors: emerging cellular biomarkers and therapeutic targets in autoimmune and inflammatory disorders","volume":"361","author":"Qamar","year":"2025","journal-title":"Life Sci."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib74","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1016\/0896-6273(90)90440-Q","article-title":"The enrichment of a neuronal growth cone collapsing activity from embryonic chick brain","volume":"4","author":"Raper","year":"1990","journal-title":"Neuron"},{"issue":"4","key":"10.1016\/j.ydbio.2026.08.012_bib75","doi-asserted-by":"crossref","first-page":"440","DOI":"10.1038\/nn2064","article-title":"Plexin-A2 and its ligand, Sema6A, control nucleus-centrosome coupling in migrating granule cells","volume":"11","author":"Renaud","year":"2008","journal-title":"Nat. Neurosci."},{"issue":"13","key":"10.1016\/j.ydbio.2026.08.012_bib76","doi-asserted-by":"crossref","DOI":"10.15252\/embj.2019102926","article-title":"Structural basis of semaphorin-plexin cis interaction","volume":"39","author":"Rozbesky","year":"2020","journal-title":"EMBO J."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib77","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.mvr.2009.10.005","article-title":"Semaphorin 5A promotes angiogenesis by increasing endothelial cell proliferation, migration, and decreasing apoptosis","volume":"79","author":"Sadanandam","year":"2010","journal-title":"Microvasc. Res."},{"issue":"3","key":"10.1016\/j.ydbio.2026.08.012_bib78","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1016\/j.semcdb.2012.11.003","article-title":"Structural insights into semaphorins and their receptors","volume":"24","author":"Siebold","year":"2013","journal-title":"Semin. Cell Dev. Biol."},{"issue":"7","key":"10.1016\/j.ydbio.2026.08.012_bib79","doi-asserted-by":"crossref","first-page":"1673","DOI":"10.1084\/jem.20072602","article-title":"Tumor angiogenesis and progression are enhanced by Sema4D produced by tumor-associated macrophages","volume":"205","author":"Sierra","year":"2008","journal-title":"J. Exp. Med."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib80","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1111\/febs.14313","article-title":"Fyn-dependent phosphorylation of PlexinA1 and PlexinA2 at conserved tyrosines is essential for zebrafish eye development","volume":"285","author":"St Clair","year":"2018","journal-title":"FEBS J."},{"issue":"21","key":"10.1016\/j.ydbio.2026.08.012_bib81","doi-asserted-by":"crossref","first-page":"3015","DOI":"10.1002\/1873-3468.13561","article-title":"PKC induces release of a functional ectodomain of the guidance cue semaphorin6A","volume":"593","author":"St Clair","year":"2019","journal-title":"FEBS Lett."},{"issue":"17","key":"10.1016\/j.ydbio.2026.08.012_bib82","doi-asserted-by":"crossref","first-page":"3923","DOI":"10.1242\/dev.01966","article-title":"Mammary ductal morphogenesis requires paracrine activation of stromal EGFR via ADAM17-dependent shedding of epithelial amphiregulin","volume":"132","author":"Sternlicht","year":"2005","journal-title":"Development"},{"issue":"15","key":"10.1016\/j.ydbio.2026.08.012_bib83","doi-asserted-by":"crossref","first-page":"10536","DOI":"10.1074\/jbc.M112.444703","article-title":"Systematic proteomic analysis identifies beta-site amyloid precursor protein cleaving enzyme 2 and 1 (BACE2 and BACE1) substrates in pancreatic beta-cells","volume":"288","author":"Stutzer","year":"2013","journal-title":"J. Biol. Chem."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib84","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1038\/s41392-024-01873-6","article-title":"Targeting a disintegrin and metalloprotease (ADAM) 17-CD122 axis enhances CD8(+) T cell effector differentiation and anti-tumor immunity","volume":"9","author":"Sun","year":"2024","journal-title":"Signal Transduct. Targeted Ther."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib85","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/S0896-6273(02)00750-X","article-title":"Plexin-B1 directly interacts with PDZ-RhoGEF\/LARG to regulate RhoA and growth cone morphology","volume":"35","author":"Swiercz","year":"2002","journal-title":"Neuron"},{"issue":"6","key":"10.1016\/j.ydbio.2026.08.012_bib86","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1083\/jcb.200312094","article-title":"Plexin-B1\/RhoGEF-mediated RhoA activation involves the receptor tyrosine kinase ErbB-2","volume":"165","author":"Swiercz","year":"2004","journal-title":"J. Cell Biol."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib87","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/S0092-8674(00)80063-X","article-title":"Plexins are a large family of receptors for transmembrane, secreted, and GPI-anchored semaphorins in vertebrates","volume":"99","author":"Tamagnone","year":"1999","journal-title":"Cell"},{"issue":"11","key":"10.1016\/j.ydbio.2026.08.012_bib88","doi-asserted-by":"crossref","DOI":"10.1002\/pro.4452","article-title":"Hybrid in vitro\/in silico analysis of low-affinity protein-protein interactions that regulate signal transduction by Sema6D","volume":"31","author":"Tanaka","year":"2022","journal-title":"Protein Sci."},{"issue":"4","key":"10.1016\/j.ydbio.2026.08.012_bib89","doi-asserted-by":"crossref","first-page":"4956","DOI":"10.1096\/fj.201902649R","article-title":"Substrate-selective protein ectodomain shedding by ADAM17 and iRhom2 depends on their juxtamembrane and transmembrane domains","volume":"34","author":"Tang","year":"2020","journal-title":"FASEB J."},{"issue":"2","key":"10.1016\/j.ydbio.2026.08.012_bib90","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1083\/jcb.151.2.209","article-title":"Ectodomain shedding of epidermal growth factor receptor ligands is required for keratinocyte migration in cutaneous wound healing","volume":"151","author":"Tokumaru","year":"2000","journal-title":"J. Cell Biol."},{"issue":"4","key":"10.1016\/j.ydbio.2026.08.012_bib91","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1101\/gad.1167304","article-title":"Dual roles of Sema6D in cardiac morphogenesis through region-specific association of its receptor, Plexin-A1, with off-track and vascular endothelial growth factor receptor type 2","volume":"18","author":"Toyofuku","year":"2004","journal-title":"Genes Dev."},{"issue":"5","key":"10.1016\/j.ydbio.2026.08.012_bib92","doi-asserted-by":"crossref","first-page":"537","DOI":"10.1016\/j.devcel.2015.11.008","article-title":"Lrig2 negatively regulates ectodomain shedding of axon guidance receptors by ADAM proteases","volume":"35","author":"van Erp","year":"2015","journal-title":"Dev. Cell"},{"issue":"20","key":"10.1016\/j.ydbio.2026.08.012_bib93","doi-asserted-by":"crossref","first-page":"14137","DOI":"10.1074\/jbc.274.20.14137","article-title":"A PDZ protein regulates the distribution of the transmembrane semaphorin, M-SemF","volume":"274","author":"Wang","year":"1999","journal-title":"J. Biol. Chem."},{"issue":"11","key":"10.1016\/j.ydbio.2026.08.012_bib94","doi-asserted-by":"crossref","first-page":"3498","DOI":"10.1182\/blood.V97.11.3498","article-title":"Functional soluble CD100\/Sema4D released from activated lymphocytes: possible role in normal and pathologic immune responses","volume":"97","author":"Wang","year":"2001","journal-title":"Blood"},{"key":"10.1016\/j.ydbio.2026.08.012_bib95","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.01279","article-title":"Structural basis for activation and non-canonical catalysis of the Rap GTPase activating protein domain of plexin","volume":"2","author":"Wang","year":"2013","journal-title":"eLife"},{"issue":"58","key":"10.1016\/j.ydbio.2026.08.012_bib96","doi-asserted-by":"crossref","first-page":"7260","DOI":"10.1038\/onc.2008.328","article-title":"Soluble monomeric EphrinA1 is released from tumor cells and is a functional ligand for the EphA2 receptor","volume":"27","author":"Wykosky","year":"2008","journal-title":"Oncogene"},{"key":"10.1016\/j.ydbio.2026.08.012_bib97","doi-asserted-by":"crossref","first-page":"2912","DOI":"10.12659\/MSM.895441","article-title":"Serum soluble semaphorin 4D is associated with left atrial diameter in patients with atrial fibrillation","volume":"21","author":"Xiang","year":"2015","journal-title":"Med. Sci. Monit. : Int. Med. J. Exp. Clin. Res."},{"issue":"1","key":"10.1016\/j.ydbio.2026.08.012_bib98","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1186\/s13075-016-1217-5","article-title":"Semaphorin 7A as a potential immune regulator and promising therapeutic target in rheumatoid arthritis","volume":"19","author":"Xie","year":"2017","journal-title":"Arthritis Res. Ther."},{"issue":"7","key":"10.1016\/j.ydbio.2026.08.012_bib99","doi-asserted-by":"crossref","first-page":"2638","DOI":"10.1523\/JNEUROSCI.20-07-02638.2000","article-title":"The transmembrane protein semaphorin 6A repels embryonic sympathetic axons","volume":"20","author":"Xu","year":"2000","journal-title":"J. Neurosci."},{"issue":"10","key":"10.1016\/j.ydbio.2026.08.012_bib100","doi-asserted-by":"crossref","first-page":"2015","DOI":"10.1158\/1541-7786.MCR-18-0933","article-title":"Semaphorin 4C promotes macrophage recruitment and angiogenesis in breast cancer","volume":"17","author":"Yang","year":"2019","journal-title":"Mol. Cancer Res."},{"issue":"4","key":"10.1016\/j.ydbio.2026.08.012_bib101","doi-asserted-by":"crossref","first-page":"230","DOI":"10.1016\/j.tins.2011.12.003","article-title":"The cis side of juxtacrine signaling: a new role in the development of the nervous system","volume":"35","author":"Yaron","year":"2012","journal-title":"Trends Neurosci."},{"issue":"10","key":"10.1016\/j.ydbio.2026.08.012_bib102","doi-asserted-by":"crossref","DOI":"10.1016\/j.celrep.2022.111785","article-title":"Tace\/ADAM17 is a bi-directional regulator of axon guidance that coordinates distinct Frazzled and Dcc receptor signaling outputs","volume":"41","author":"Zang","year":"2022","journal-title":"Cell Rep."},{"issue":"9","key":"10.1016\/j.ydbio.2026.08.012_bib103","doi-asserted-by":"crossref","first-page":"3537","DOI":"10.1021\/pr070247w","article-title":"A comparative proteomic analysis of human and rat embryonic cerebrospinal fluid","volume":"6","author":"Zappaterra","year":"2007","journal-title":"J. Proteome Res."},{"issue":"9","key":"10.1016\/j.ydbio.2026.08.012_bib104","first-page":"16352","article-title":"Serum Sema4D levels are associated with lumbar spine bone mineral density and bone turnover markers in patients with postmenopausal osteoporosis","volume":"8","author":"Zhang","year":"2015","journal-title":"Int. J. Clin. Exp. Med."},{"issue":"2","key":"10.1016\/j.ydbio.2026.08.012_bib105","doi-asserted-by":"crossref","first-page":"456","DOI":"10.1016\/j.celrep.2017.12.044","article-title":"PlexinA2 forward signaling through Rap1 GTPases regulates dentate gyrus development and Schizophrenia-like behaviors","volume":"22","author":"Zhao","year":"2018","journal-title":"Cell Rep."},{"issue":"21","key":"10.1016\/j.ydbio.2026.08.012_bib106","doi-asserted-by":"crossref","DOI":"10.3390\/ijms232112796","article-title":"Identification of molecular determinants in iRhoms1 and 2 That contribute to the substrate selectivity of stimulated ADAM17","volume":"23","author":"Zhao","year":"2022","journal-title":"Int. J. Mol. Sci."},{"issue":"5","key":"10.1016\/j.ydbio.2026.08.012_bib107","doi-asserted-by":"crossref","first-page":"1621","DOI":"10.1073\/pnas.0606344104","article-title":"Regulated surface expression and shedding support a dual role for semaphorin 4D in platelet responses to vascular injury","volume":"104","author":"Zhu","year":"2007","journal-title":"Proc. Natl. Acad. Sci. U. S. A."}],"container-title":["Developmental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0012160626001855?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0012160626001855?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T07:57:00Z","timestamp":1790582220000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0012160626001855"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,11]]},"references-count":107,"alternative-id":["S0012160626001855"],"URL":"https:\/\/doi.org\/10.1016\/j.ydbio.2026.08.012","relation":{},"ISSN":["0012-1606"],"issn-type":[{"value":"0012-1606","type":"print"}],"subject":[],"published":{"date-parts":[[2026,11]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Shedding light on the proteolytically released ectodomains of membrane-bound Semaphorins","name":"articletitle","label":"Article Title"},{"value":"Developmental Biology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.ydbio.2026.08.012","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 Elsevier Inc. All rights are reserved, including those for text and data mining, AI training, and similar technologies.","name":"copyright","label":"Copyright"}]}}