{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,19]],"date-time":"2026-09-19T09:09:12Z","timestamp":1789808952221,"version":"4.0.1"},"reference-count":71,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,8,15]],"date-time":"2025-08-15T00:00:00Z","timestamp":1755216000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,8,15]],"date-time":"2025-08-15T00:00:00Z","timestamp":1755216000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","award":["ANR-PRCI-MOTAC80C\/A134\/AN16HRJNMF"],"award-info":[{"award-number":["ANR-PRCI-MOTAC80C\/A134\/AN16HRJNMF"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","award":["ANR-PRC-MOTOX80C\/213Z\/AN20HRJNMF"],"award-info":[{"award-number":["ANR-PRC-MOTOX80C\/213Z\/AN20HRJNMF"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","award":["ANR-PRC-SYMPA80C\/213Z\/AN21HRJNMF"],"award-info":[{"award-number":["ANR-PRC-SYMPA80C\/213Z\/AN21HRJNMF"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","award":["ANR-PRC-MOTOX80C\/213Z\/AN20HRJNMF"],"award-info":[{"award-number":["ANR-PRC-MOTOX80C\/213Z\/AN20HRJNMF"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","award":["ANR-PRC-SYMPA80C\/213Z\/AN21HRJNMF"],"award-info":[{"award-number":["ANR-PRC-SYMPA80C\/213Z\/AN21HRJNMF"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Commun Biol"],"abstract":"<jats:title>Abstract<\/jats:title>\n          <jats:p>Cerebrospinal Fluid-contacting neurons (CSF-cNs) are GABAergic bipolar neurons found, in contact with the cerebrospinal fluid, along the vertebrate medullo-spinal central canal. They express Polycystin Kidney Disease 2-Like 1 channels (PKD2L1), members of the Transient Receptor Potential superfamily, and were shown to modulate motor activity and therefore suggested to act as a novel sensory system. However, in mice, they remain largely uncharacterized and it is crucial to comprehensively characterize their morphological and electrophysiological properties to determine whether they form a homogenous neuronal population and understand their role in the CNS. We show that CSF-cNs are distributed throughout the spinal cord with a uniform morphology and a primarily ventral localization. They exhibit region-specific properties, expression of voltage-dependent and ligand-gated conductances and detect variation in extracellular pH through activation of PKD2L1 and Acid-sensing Ion Channels. They possess GABA<jats:sub>B<\/jats:sub> and muscarinic receptors, not glutamatergic metabotropic ones, to modulate Ca<jats:sup>2+<\/jats:sup> channels. CSF-cNs represent unique sensory neurons with a uniform morphology and electrophysiological properties that appear specific to the spinal cord segment inserted in. The future challenges in the field, will be to elucidate the physiological stimuli activating CSF-cNs and the neuronal network they are integrated in to modulate body function through specific local spinal network.<\/jats:p>","DOI":"10.1038\/s42003-025-08559-x","type":"journal-article","created":{"date-parts":[[2025,8,15]],"date-time":"2025-08-15T17:53:57Z","timestamp":1755280437000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["Cerebrospinal fluid-contacting neurons are sensory neurons with uniform morphological and region-specific electrophysiological properties in the mouse spinal cord"],"prefix":"10.1038","volume":"8","author":[{"ORCID":"https:\/\/orcid.org\/0009-0005-7440-3966","authenticated-orcid":false,"given":"Elysa","family":"Crozat","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-4378-1697","authenticated-orcid":false,"given":"Edith","family":"Blasco","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2426-6140","authenticated-orcid":false,"given":"Jorge","family":"Ramirez-Franco","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Priscille","family":"Riondel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5076-9753","authenticated-orcid":false,"given":"Nina","family":"Jur\u010di\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5052-4767","authenticated-orcid":false,"given":"Riad","family":"Seddik","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9235-074X","authenticated-orcid":false,"given":"Caroline","family":"Michelle","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4235-7281","authenticated-orcid":false,"given":"J\u00e9r\u00f4me","family":"Trouslard","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5486-2771","authenticated-orcid":false,"given":"Nicolas","family":"Wanaverbecq","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,8,15]]},"reference":[{"key":"8559_CR1","doi-asserted-by":"publisher","first-page":"236","DOI":"10.1007\/BF00406356","volume":"13","author":"W Kolmer","year":"1931","unstructured":"Kolmer, W. \u00dcber das Sagittalorgan, ein zentrales Sinnesorgan der Wirbeltiere, insbesondere beim Affen. Z. Zellforsch. Mikrosk. Anat. 13, 236\u2013248 (1931).","journal-title":"Z. Zellforsch. Mikrosk. Anat."},{"key":"8559_CR2","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1007\/BF02593586","volume":"66","author":"E Agduhr","year":"1922","unstructured":"Agduhr, E. \u00dcber ein zentrales Sinnesorgan (?) bei den Vertebraten. Z. Anat. Entwicklungsgeschichte 66, 223\u2013360 (1922).","journal-title":"Z. Anat. Entwicklungsgeschichte"},{"key":"8559_CR3","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1002\/(SICI)1097-0029(19980401)41:1<57::AID-JEMT6>3.0.CO;2-R","volume":"41","author":"B Vigh","year":"1998","unstructured":"Vigh, B. & Vigh-Teichmann, I. Actual problems of the cerebrospinal fluid-contacting neurons. Microsc. Res. Tech. 41, 57\u201383 (1998).","journal-title":"Microsc. Res. Tech."},{"key":"8559_CR4","doi-asserted-by":"publisher","first-page":"527","DOI":"10.1002\/cne.902580405","volume":"258","author":"CC LaMotte","year":"1987","unstructured":"LaMotte, C. C. Vasoactive intestinal polypeptide cerebrospinal fluid-contacting neurons of the monkey and cat spinal central canal. J. Comp. Neurol. 258, 527\u2013541 (1987).","journal-title":"J. Comp. Neurol."},{"key":"8559_CR5","doi-asserted-by":"publisher","first-page":"26","DOI":"10.3389\/fnana.2014.00026","volume":"8","author":"L Djenoune","year":"2014","unstructured":"Djenoune, L. et al. Investigation of spinal cerebrospinal fluid-contacting neurons expressing PKD2L1: evidence for a conserved system from fish to primates. Front Neuroanat. 8, 26 (2014).","journal-title":"Front Neuroanat."},{"key":"8559_CR6","doi-asserted-by":"publisher","first-page":"1753","DOI":"10.1002\/cne.23542","volume":"522","author":"E Jalalvand","year":"2014","unstructured":"Jalalvand, E., Robertson, B., Wall\u00e9n, P., Hill, R. H. & Grillner, S. Laterally projecting cerebrospinal fluid-contacting cells in the lamprey spinal cord are of two distinct types. J. Comp. Neurol. 522, 1753\u20131768 (2014).","journal-title":"J. Comp. Neurol."},{"key":"8559_CR7","doi-asserted-by":"publisher","first-page":"e87748","DOI":"10.1371\/journal.pone.0087748","volume":"9","author":"A Orts-Del\u2019Immagine","year":"2014","unstructured":"Orts-Del\u2019Immagine, A. et al. Morphology, distribution and phenotype of polycystin kidney disease 2-like 1-positive cerebrospinal fluid contacting neurons in the brainstem of adult mice. PLoS ONE 9, e87748 (2014).","journal-title":"PLoS ONE"},{"key":"8559_CR8","first-page":"880","volume":"143","author":"YL Petracca","year":"2016","unstructured":"Petracca, Y. L. et al. The late and dual origin of cerebrospinal fluid-contacting neurons in the mouse spinal cord. Development 143, 880\u2013891 (2016).","journal-title":"Development"},{"key":"8559_CR9","doi-asserted-by":"publisher","first-page":"10010","DOI":"10.1523\/JNEUROSCI.6183-08.2009","volume":"29","author":"N Marichal","year":"2009","unstructured":"Marichal, N., Garc\u00eda, G., Radmilovich, M., Trujillo-Cen\u00f3z, O. & Russo, R. E. Enigmatic central canal contacting cells: immature neurons in \u2018standby mode\u2019?. J. Neurosci. 29, 10010\u201310024 (2009).","journal-title":"J. Neurosci."},{"key":"8559_CR10","doi-asserted-by":"publisher","first-page":"13582","DOI":"10.3390\/ijms241713582","volume":"24","author":"X Liu","year":"2023","unstructured":"Liu, X. et al. A Comparison of PKD2L1-Expressing Cerebrospinal Fluid Contacting Neurons in Spinal Cords of Rodents, Carnivores, and Primates. Int. J. Mol. Sci. 24, 13582 (2023).","journal-title":"Int. J. Mol. Sci."},{"key":"8559_CR11","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1002\/cne.10565","volume":"457","author":"M-E Stoeckel","year":"2003","unstructured":"Stoeckel, M.-E. et al. Cerebrospinal fluid-contacting neurons in the rat spinal cord, a gamma-aminobutyric acidergic system expressing the P2X2 subunit of purinergic receptors, PSA-NCAM, and GAP-43 immunoreactivities: light and electron microscopic study. J. Comp. Neurol. 457, 159\u2013174 (2003).","journal-title":"J. Comp. Neurol."},{"key":"8559_CR12","doi-asserted-by":"publisher","first-page":"3719","DOI":"10.1113\/jphysiol.2012.227959","volume":"590","author":"A Orts-Del\u2019Immagine","year":"2012","unstructured":"Orts-Del\u2019Immagine, A. et al. Properties of subependymal cerebrospinal fluid contacting neurones in the dorsal vagal complex of the mouse brainstem. J. Physiol. (Lond.) 590, 3719\u20133741 (2012).","journal-title":"J. Physiol. (Lond.)"},{"key":"8559_CR13","doi-asserted-by":"crossref","unstructured":"Jur\u010di\u0107, N., Michelle, C., Trouslard, J., Wanaverbecq, N. & Kastner, A. Evidence for PKD2L1-positive neurons distant from the central canal in the ventromedial spinal cord and medulla of the adult mouse. Eur. J. Neurosci. 54, 4781\u20134803 (2021).","DOI":"10.1111\/ejn.15342"},{"key":"8559_CR14","doi-asserted-by":"publisher","first-page":"e83108","DOI":"10.7554\/eLife.83108","volume":"12","author":"Y Nakamura","year":"2023","unstructured":"Nakamura, Y. et al. Cerebrospinal fluid-contacting neuron tracing reveals structural and functional connectivity for locomotion in the mouse spinal cord. Elife 12, e83108 (2023).","journal-title":"Elife"},{"key":"8559_CR15","doi-asserted-by":"publisher","first-page":"2442","DOI":"10.1016\/j.cub.2022.04.019","volume":"32","author":"K Gerstmann","year":"2022","unstructured":"Gerstmann, K. et al. The role of intraspinal sensory neurons in the control of quadrupedal locomotion. Curr. Biol. 32, 2442\u20132453.e4 (2022).","journal-title":"Curr. Biol."},{"key":"8559_CR16","doi-asserted-by":"publisher","first-page":"407","DOI":"10.1038\/nature08323","volume":"461","author":"C Wyart","year":"2009","unstructured":"Wyart, C. et al. Optogenetic dissection of a behavioural module in the vertebrate spinal cord. Nature 461, 407\u2013410 (2009).","journal-title":"Nature"},{"key":"8559_CR17","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-017-00350-1","volume":"7","author":"L Djenoune","year":"2017","unstructured":"Djenoune, L. et al. The dual developmental origin of spinal cerebrospinal fluid-contacting neurons gives rise to distinct functional subtypes. Sci. Rep. 7, 719 (2017).","journal-title":"Sci. Rep."},{"key":"8559_CR18","doi-asserted-by":"publisher","first-page":"631","DOI":"10.1113\/JP277172","volume":"597","author":"N Jur\u010di\u0107","year":"2019","unstructured":"Jur\u010di\u0107, N. et al. GABAB receptors modulate Ca2+ but not G protein-gated inwardly rectifying K+ channels in cerebrospinal-fluid contacting neurones of mouse brainstem. J. Physiol. (Lond.) 597, 631\u2013651 (2019).","journal-title":"J. Physiol. (Lond.)"},{"key":"8559_CR19","doi-asserted-by":"publisher","first-page":"105914","DOI":"10.1016\/j.isci.2022.105914","volume":"26","author":"E Johnson","year":"2023","unstructured":"Johnson, E. et al. Graded spikes differentially signal neurotransmitter input in cerebrospinal fluid contacting neurons of the mouse spinal cord. iScience 26, 105914 (2023).","journal-title":"iScience"},{"key":"8559_CR20","doi-asserted-by":"publisher","first-page":"176","DOI":"10.1126\/science.aam8999","volume":"360","author":"AB Rosenberg","year":"2018","unstructured":"Rosenberg, A. B. et al. Single-cell profiling of the developing mouse brain and spinal cord with split-pool barcoding. Science 360, 176\u2013182 (2018).","journal-title":"Science"},{"key":"8559_CR21","doi-asserted-by":"publisher","first-page":"549","DOI":"10.1016\/j.neuropharm.2015.07.030","volume":"101","author":"A Orts-Del\u2019Immagine","year":"2016","unstructured":"Orts-Del\u2019Immagine, A. et al. A single polycystic kidney disease 2-like 1 channel opening acts as a spike generator in cerebrospinal fluid-contacting neurons of adult mouse brainstem. Neuropharmacology 101, 549\u2013565 (2016).","journal-title":"Neuropharmacology"},{"key":"8559_CR22","doi-asserted-by":"publisher","first-page":"1346","DOI":"10.1016\/j.cub.2016.03.048","volume":"26","author":"E Jalalvand","year":"2016","unstructured":"Jalalvand, E., Robertson, B., Tostivint, H., Wall\u00e9n, P. & Grillner, S. The Spinal Cord Has an Intrinsic System for the Control of pH. Curr. Biol. 26, 1346\u20131351 (2016).","journal-title":"Curr. Biol."},{"key":"8559_CR23","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms10002","volume":"7","author":"E Jalalvand","year":"2016","unstructured":"Jalalvand, E., Robertson, B., Wall\u00e9n, P. & Grillner, S. Ciliated neurons lining the central canal sense both fluid movement and pH through ASIC3. Nat. Commun. 7, 10002 (2016).","journal-title":"Nat. Commun."},{"key":"8559_CR24","doi-asserted-by":"publisher","first-page":"934","DOI":"10.1038\/nature05084","volume":"442","author":"AL Huang","year":"2006","unstructured":"Huang, A. L. et al. The cells and logic for mammalian sour taste detection. Nature 442, 934\u2013938 (2006).","journal-title":"Nature"},{"key":"8559_CR25","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms10866","volume":"7","author":"UL B\u00f6hm","year":"2016","unstructured":"B\u00f6hm, U. L. et al. CSF-contacting neurons regulate locomotion by relaying mechanical stimuli to spinal circuits. Nat. Commun. 7, 10866 (2016).","journal-title":"Nat. Commun."},{"key":"8559_CR26","doi-asserted-by":"publisher","first-page":"827","DOI":"10.1016\/j.cub.2019.12.071","volume":"30","author":"A Orts-Del\u2019Immagine","year":"2020","unstructured":"Orts-Del\u2019Immagine, A. et al. Sensory Neurons Contacting the Cerebrospinal Fluid Require the Reissner Fiber to Detect Spinal Curvature In Vivo. Curr. Biol. 30, 827\u2013839.e4 (2020).","journal-title":"Curr. Biol."},{"key":"8559_CR27","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-018-06225-x","volume":"9","author":"JR Sternberg","year":"2018","unstructured":"Sternberg, J. R. et al. Pkd2l1 is required for mechanoception in cerebrospinal fluid-contacting neurons and maintenance of spine curvature. Nat. Commun. 9, 3804 (2018).","journal-title":"Nat. Commun."},{"key":"8559_CR28","doi-asserted-by":"publisher","first-page":"2264","DOI":"10.1016\/j.celrep.2019.07.087","volume":"28","author":"DJ Di Bella","year":"2019","unstructured":"Di Bella, D. J. et al. Ascl1 Balances Neuronal versus Ependymal Fate in the Spinal Cord Central Canal. Cell Rep. 28, 2264\u20132274.e3 (2019).","journal-title":"Cell Rep."},{"key":"8559_CR29","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1016\/j.neuroscience.2016.11.028","volume":"343","author":"A Orts-Del\u2019Immagine","year":"2017","unstructured":"Orts-Del\u2019Immagine, A. et al. Postnatal maturation of mouse medullo-spinal cerebrospinal fluid-contacting neurons. Neuroscience 343, 39\u201354 (2017).","journal-title":"Neuroscience"},{"key":"8559_CR30","doi-asserted-by":"publisher","first-page":"190","DOI":"10.1002\/(SICI)1097-4695(199810)37:1<190::AID-NEU14>3.0.CO;2-I","volume":"37","author":"NC Spitzer","year":"1998","unstructured":"Spitzer, N. C. & Ribera, A. B. Development of electrical excitability in embryonic neurons: mechanisms and roles. J. Neurobiol. 37, 190\u2013197 (1998).","journal-title":"J. Neurobiol."},{"key":"8559_CR31","doi-asserted-by":"publisher","first-page":"e2289222024","DOI":"10.1523\/JNEUROSCI.2289-22.2024","volume":"44","author":"P Riondel","year":"2024","unstructured":"Riondel, P. et al. Evidence for Two Subpopulations of Cerebrospinal Fluid-Contacting Neurons with Opposite GABAergic Signaling in Adult Mouse Spinal Cord. J. Neurosci. 44, e2289222024 (2024).","journal-title":"J. Neurosci."},{"key":"8559_CR32","doi-asserted-by":"publisher","first-page":"103","DOI":"10.1016\/j.conb.2013.12.016","volume":"26","author":"K Fidelin","year":"2014","unstructured":"Fidelin, K. & Wyart, C. Inhibition and motor control in the developing zebrafish spinal cord. Curr. Opin. Neurobiol. 26, 103\u2013109 (2014).","journal-title":"Curr. Opin. Neurobiol."},{"key":"8559_CR33","doi-asserted-by":"publisher","first-page":"2841","DOI":"10.1016\/j.cub.2016.08.026","volume":"26","author":"JM Hubbard","year":"2016","unstructured":"Hubbard, J. M. et al. Intraspinal Sensory Neurons Provide Powerful Inhibition to Motor Circuits Ensuring Postural Control during Locomotion. Curr. Biol. 26, 2841\u20132853 (2016).","journal-title":"Curr. Biol."},{"key":"8559_CR34","doi-asserted-by":"publisher","first-page":"940","DOI":"10.1016\/j.cub.2023.01.039","volume":"33","author":"AE Prendergast","year":"2023","unstructured":"Prendergast, A. E. et al. CSF-contacting neurons respond to Streptococcus pneumoniae and promote host survival during central nervous system infection. Curr. Biol. 33, 940\u2013956.e10 (2023).","journal-title":"Curr. Biol."},{"key":"8559_CR35","doi-asserted-by":"publisher","unstructured":"Yue, W. W. S., Touhara, K. K., Toma, K., Duan, X. & Julius, D. Endogenous opioid signalling regulates spinal ependymal cell proliferation. Nature 1\u20138 https:\/\/doi.org\/10.1038\/s41586-024-07889-w (2024).","DOI":"10.1038\/s41586-024-07889-w"},{"key":"8559_CR36","doi-asserted-by":"publisher","first-page":"5633","DOI":"10.1113\/jphysiol.2011.214312","volume":"589","author":"C Reali","year":"2011","unstructured":"Reali, C., Fern\u00e1ndez, A., Radmilovich, M., Trujillo-Cen\u00f3z, O. & Russo, R. E. GABAergic signalling in a neurogenic niche of the turtle spinal cord. J. Physiol. 589, 5633\u20135647 (2011).","journal-title":"J. Physiol."},{"key":"8559_CR37","doi-asserted-by":"publisher","unstructured":"Kastner, A. & Wanaverbecq, N. In Rhesus monkeys, CSF-contacting neurons are the only neurons present in the medullo-spinal peri-ependymal zone. 2023.03.29.534787 Preprint at https:\/\/doi.org\/10.1101\/2023.03.29.534787 (2023).","DOI":"10.1101\/2023.03.29.534787"},{"key":"8559_CR38","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1038\/s41593-018-0301-3","volume":"22","author":"R Cai","year":"2019","unstructured":"Cai, R. et al. Panoptic imaging of transparent mice reveals whole-body neuronal projections and skull-meninges connections. Nat. Neurosci. 22, 317\u2013327 (2019).","journal-title":"Nat. Neurosci."},{"key":"8559_CR39","doi-asserted-by":"publisher","first-page":"1197","DOI":"10.1038\/s41596-022-00788-2","volume":"18","author":"R Cai","year":"2023","unstructured":"Cai, R. et al. Whole-mouse clearing and imaging at the cellular level with vDISCO. Nat. Protoc. 18, 1197\u20131242 (2023).","journal-title":"Nat. Protoc."},{"key":"8559_CR40","doi-asserted-by":"publisher","first-page":"2864","DOI":"10.1002\/stem.2077","volume":"33","author":"LF Corns","year":"2015","unstructured":"Corns, L. F. et al. Cholinergic Enhancement of Cell Proliferation in the Postnatal Neurogenic Niche of the Mammalian Spinal Cord. Stem Cells 33, 2864\u20132876 (2015).","journal-title":"Stem Cells"},{"key":"8559_CR41","doi-asserted-by":"publisher","unstructured":"Kutna, V., Sevc, J., Gombalov\u00e1, Z., Matia\u0161ov\u00e1, A. & Daxnerov\u00e1, Z. Enigmatic cerebrospinal fluid-contacting neurons arise even after the termination of neurogenesis in the rat spinal cord during embryonic development and retain their immature-like characteristics until adulthood. Acta Histochem. https:\/\/doi.org\/10.1016\/j.acthis.2013.08.004 (2013).","DOI":"10.1016\/j.acthis.2013.08.004"},{"key":"8559_CR42","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/0165-0270(88)90021-0","volume":"23","author":"R Bjugn","year":"1988","unstructured":"Bjugn, R., Haugland, H. K. & Flood, P. R. Preservation of the ependyma of the mouse spinal cord by various vascular perfusion procedures. J. Neurosci. Methods 23, 49\u201355 (1988).","journal-title":"J. Neurosci. Methods"},{"key":"8559_CR43","doi-asserted-by":"publisher","unstructured":"Tonelli Gombalov\u00e1, Z. et al. Majority of cerebrospinal fluid-contacting neurons in the spinal cord of C57Bl\/6\u2009N mice is present in ectopic position unlike in other studied experimental mice strains and mammalian species. J. Comp. Neurol. https:\/\/doi.org\/10.1002\/cne.24909 (2020).","DOI":"10.1002\/cne.24909"},{"key":"8559_CR44","doi-asserted-by":"publisher","first-page":"2235","DOI":"10.3390\/cells10092235","volume":"10","author":"A Torrillas de la Cal","year":"2021","unstructured":"Torrillas de la Cal, A. et al. The Structure of the Spinal Cord Ependymal Region in Adult Humans Is a Distinctive Trait among Mammals. Cells 10, 2235 (2021).","journal-title":"Cells"},{"key":"8559_CR45","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1002\/ca.24196","volume":"38","author":"E Lefevre","year":"2025","unstructured":"Lefevre, E. et al. Upper end of the central canal of the human spinal cord: Quantitative anatomical study and 3D modeling. Clin. Anat. 38, 35\u201342 (2025).","journal-title":"Clin. Anat."},{"key":"8559_CR46","doi-asserted-by":"crossref","unstructured":"Saker, E. et al. The Human Central Canal of the Spinal Cord: A Comprehensive Review of its Anatomy, Embryology, Molecular Development, Variants, and Pathology. Cureus 8, e927 (2016).","DOI":"10.7759\/cureus.927"},{"key":"8559_CR47","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1007\/s004010050982","volume":"97","author":"K Yasui","year":"1999","unstructured":"Yasui, K., Hashizume, Y., Yoshida, M., Kameyama, T. & Sobue, G. Age-related morphologic changes of the central canal of the human spinal cord. Acta Neuropathol. 97, 253\u2013259 (1999).","journal-title":"Acta Neuropathol."},{"key":"8559_CR48","doi-asserted-by":"publisher","first-page":"328","DOI":"10.1016\/j.neuron.2023.01.007","volume":"111","author":"A Yadav","year":"2023","unstructured":"Yadav, A. et al. A cellular taxonomy of the adult human spinal cord. Neuron 111, 328\u2013344.e7 (2023).","journal-title":"Neuron"},{"key":"8559_CR49","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1002\/cne.900860102","volume":"86","author":"T Humphrey","year":"1947","unstructured":"Humphrey, T. Sensory ganglion cells within the central canal of the embryonic human spinal cord. J. Comp. Neurol. 86, 1\u201335 (1947).","journal-title":"J. Comp. Neurol."},{"key":"8559_CR50","doi-asserted-by":"crossref","unstructured":"Urrutia-Pi\u00f1ones, J., Morales-Moraga, C., Sanguinetti-Gonz\u00e1lez, N., Escobar, A. P. & Chiu, C. Q. Long-Range GABAergic Projections of Cortical Origin in Brain Function. Front. Syst. Neurosci. 16, 841869 (2022).","DOI":"10.3389\/fnsys.2022.841869"},{"key":"8559_CR51","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1038\/336170a0","volume":"336","author":"TF Freund","year":"1988","unstructured":"Freund, T. F. & Antal, M. GABA-containing neurons in the septum control inhibitory interneurons in the hippocampus. Nature 336, 170\u2013173 (1988).","journal-title":"Nature"},{"key":"8559_CR52","doi-asserted-by":"publisher","first-page":"391","DOI":"10.1016\/S0166-2236(98)01310-1","volume":"21","author":"CI De Zeeuw","year":"1998","unstructured":"De Zeeuw, C. I. et al. Microcircuitry and function of the inferior olive. Trends Neurosci. 21, 391\u2013400 (1998).","journal-title":"Trends Neurosci."},{"key":"8559_CR53","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.cub.2015.09.070","volume":"25","author":"K Fidelin","year":"2015","unstructured":"Fidelin, K. et al. State-Dependent Modulation of Locomotion by GABAergic Spinal Sensory Neurons. Curr. Biol. 25, 1\u201313 (2015).","journal-title":"Curr. Biol."},{"key":"8559_CR54","doi-asserted-by":"publisher","first-page":"1063","DOI":"10.1523\/JNEUROSCI.3740-04.2005","volume":"25","author":"SA Deuchars","year":"2005","unstructured":"Deuchars, S. A., Milligan, C. J., Stornetta, R. L. & Deuchars, J. GABAergic neurons in the central region of the spinal cord: a novel substrate for sympathetic inhibition. J. Neurosci. 25, 1063\u20131070 (2005).","journal-title":"J. Neurosci."},{"key":"8559_CR55","doi-asserted-by":"crossref","unstructured":"Orts-Del'Immagine, A. et al. A single polycystic kidney disease 2-like 1 channel opening acts as a spike generator in cerebrospinal fluid-contacting neurons of adult mouse brainstem. Neuropharmacology 101, 549\u2013565 (2016).","DOI":"10.1016\/j.neuropharm.2015.07.030"},{"key":"8559_CR56","doi-asserted-by":"publisher","first-page":"485","DOI":"10.1016\/j.tins.2010.08.005","volume":"33","author":"Y Ben-Ari","year":"2010","unstructured":"Ben-Ari, Y. & Spitzer, N. C. Phenotypic checkpoints regulate neuronal development. Trends Neurosci. 33, 485\u2013492 (2010).","journal-title":"Trends Neurosci."},{"key":"8559_CR57","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1038\/380255a0","volume":"380","author":"SR Ikeda","year":"1996","unstructured":"Ikeda, S. R. Voltage-dependent modulation of N-type calcium channels by G-protein beta gamma subunits. Nature 380, 255\u2013258 (1996).","journal-title":"Nature"},{"key":"8559_CR58","doi-asserted-by":"publisher","first-page":"652","DOI":"10.1007\/BF02593657","volume":"60","author":"WDas Kolmer","year":"1921","unstructured":"Kolmer, W. D. as \u201eSagittalorgan\u201c der Wirbeltiere. Z. f\u00fcr Anat. und Entwicklungsgeschichte 60, 652\u2013717 (1921).","journal-title":"Z. f\u00fcr Anat. und Entwicklungsgeschichte"},{"key":"8559_CR59","doi-asserted-by":"publisher","first-page":"796","DOI":"10.1016\/j.cell.2020.01.030","volume":"180","author":"S Zhao","year":"2020","unstructured":"Zhao, S. et al. Cellular and Molecular Probing of Intact Human Organs. Cell 180, 796\u2013812.e19 (2020).","journal-title":"Cell"},{"key":"8559_CR60","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","volume":"9","author":"J Schindelin","year":"2012","unstructured":"Schindelin, J. et al. Fiji: an open-source platform for biological-image analysis. Nat. Methods 9, 676\u2013682 (2012).","journal-title":"Nat. Methods"},{"key":"8559_CR61","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1002\/pro.3235","volume":"27","author":"TD Goddard","year":"2018","unstructured":"Goddard, T. D. et al. UCSF ChimeraX: Meeting modern challenges in visualization and analysis. Protein Sci. 27, 14\u201325 (2018).","journal-title":"Protein Sci."},{"key":"8559_CR62","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1002\/pro.3943","volume":"30","author":"EF Pettersen","year":"2021","unstructured":"Pettersen, E. F. et al. UCSF ChimeraX: Structure visualization for researchers, educators, and developers. Protein Sci. 30, 70\u201382 (2021).","journal-title":"Protein Sci."},{"key":"8559_CR63","doi-asserted-by":"publisher","first-page":"870","DOI":"10.1038\/s41592-019-0501-0","volume":"16","author":"D H\u00f6rl","year":"2019","unstructured":"H\u00f6rl, D. et al. BigStitcher: reconstructing high-resolution image datasets of cleared and expanded samples. Nat. Methods 16, 870\u2013874 (2019).","journal-title":"Nat. Methods"},{"key":"8559_CR64","doi-asserted-by":"crossref","unstructured":"Sternberg. Biomedical Image Processing. Computer 16, 22\u201334 (1983).","DOI":"10.1109\/MC.1983.1654163"},{"key":"8559_CR65","doi-asserted-by":"publisher","first-page":"1463","DOI":"10.1093\/bioinformatics\/btp184","volume":"25","author":"S Preibisch","year":"2009","unstructured":"Preibisch, S., Saalfeld, S. & Tomancak, P. Globally optimal stitching of tiled 3D microscopic image acquisitions. Bioinformatics 25, 1463\u20131465 (2009).","journal-title":"Bioinformatics"},{"key":"8559_CR66","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1038\/s41592-019-0650-1","volume":"17","author":"R Haase","year":"2020","unstructured":"Haase, R. et al. CLIJ: GPU-accelerated image processing for everyone. Nat. Methods 17, 5\u20136 (2020).","journal-title":"Nat. Methods"},{"key":"8559_CR67","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1111\/j.1365-2818.2006.01706.x","volume":"224","author":"S Bolte","year":"2006","unstructured":"Bolte, S. & Cordeli\u00e8res, F. P. A guided tour into subcellular colocalization analysis in light microscopy. J. Microsc 224, 213\u2013232 (2006).","journal-title":"J. Microsc"},{"key":"8559_CR68","doi-asserted-by":"publisher","first-page":"1840","DOI":"10.1093\/bioinformatics\/btt276","volume":"29","author":"J Ollion","year":"2013","unstructured":"Ollion, J., Cochennec, J., Loll, F., Escud\u00e9, C. & Boudier, T. TANGO: a generic tool for high-throughput 3D image analysis for studying nuclear organization. Bioinformatics 29, 1840\u20131841 (2013).","journal-title":"Bioinformatics"},{"key":"8559_CR69","doi-asserted-by":"publisher","first-page":"368","DOI":"10.1006\/cviu.1996.0065","volume":"64","author":"G Borgefors","year":"1996","unstructured":"Borgefors, G. On Digital Distance Transforms in Three Dimensions. Computer Vis. Image Underst. 64, 368\u2013376 (1996).","journal-title":"Computer Vis. Image Underst."},{"key":"8559_CR70","doi-asserted-by":"publisher","unstructured":"Linear Mixed-Effects Models: Basic Concepts and Examples. in Mixed-Effects Models in S and S-PLUS (eds. Pinheiro, J. C. & Bates, D. M.) 3\u201356 (Springer, New York, NY, 2000). https:\/\/doi.org\/10.1007\/0-387-22747-4_1.","DOI":"10.1007\/0-387-22747-4_1"},{"key":"8559_CR71","doi-asserted-by":"publisher","unstructured":"Crozat, E. et al. Spinal CSF-contacting neurons properties. https:\/\/doi.org\/10.6084\/m9.figshare.29430869 (2025).","DOI":"10.6084\/m9.figshare.29430869"}],"container-title":["Communications Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-08559-x.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-08559-x","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-08559-x.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,9,9]],"date-time":"2025-09-09T04:56:27Z","timestamp":1757393787000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s42003-025-08559-x"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,15]]},"references-count":71,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["8559"],"URL":"https:\/\/doi.org\/10.1038\/s42003-025-08559-x","relation":{},"ISSN":["2399-3642"],"issn-type":[{"value":"2399-3642","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,15]]},"assertion":[{"value":"22 November 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"17 July 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"15 August 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"1233"}}