{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T22:51:57Z","timestamp":1787698317829,"version":"build-2784847793"},"reference-count":93,"publisher":"Society for Neuroscience","issue":"14","license":[{"start":{"date-parts":[[2017,10,5]],"date-time":"2017-10-05T00:00:00Z","timestamp":1507161600000},"content-version":"vor","delay-in-days":211,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["J. Neurosci."],"published-print":{"date-parts":[[2017,4,5]]},"abstract":"<jats:p>Axonal degeneration is a pathophysiological mechanism common to several neurodegenerative diseases. The slow Wallerian degeneration (Wld<jats:sup>S<\/jats:sup>) mutation, which results in reduced axonal degeneration in the central and peripheral nervous systems, has provided insight into a redox-dependent mechanism by which axons undergo self-destruction. We studied early molecular events in axonal degeneration with single-axon laser axotomy and time-lapse imaging, monitoring the initial changes in transected axons of purified retinal ganglion cells (RGCs) from wild-type and Wld<jats:sup>S<\/jats:sup>rat retinas using a polarity-sensitive annexin-based biosensor (annexin B12-Cys101,Cys260-N,N\u2032-dimethyl-N-(iodoacetyl)-N\u2032-(7-nitrobenz-2-oxa-1,3-diazol-4-yl) ethylenediamine). Transected axons demonstrated a rapid and progressive change in membrane phospholipid polarity, manifested as phosphatidylserine externalization, which was significantly delayed and propagated more slowly in axotomized Wld<jats:sup>S<\/jats:sup>RGCs compared with wild-type axons. Delivery of bis(3-propionic acid methyl ester)phenylphosphine borane complex, a cell-permeable intracellular disulfide-reducing drug, slowed the onset and velocity of phosphatidylserine externalization in wild-type axons significantly, replicating the Wld<jats:sup>S<\/jats:sup>phenotype, whereas extracellular redox modulation reversed the Wld<jats:sup>S<\/jats:sup>phenotype. These findings are consistent with an intra-axonal redox mechanism for axonal degeneration associated with the initiation and propagation of phosphatidylserine externalization after axotomy.<\/jats:p><jats:p><jats:bold>SIGNIFICANCE STATEMENT<\/jats:bold>Axonal degeneration is a neuronal process independent of somal apoptosis, the propagation of which is unclear. We combined single-cell laser axotomy with time-lapse imaging to study the dynamics of phosphatidylserine externalization immediately after axonal injury in purified retinal ganglion cells. The extension of phosphatidylserine externalization was slowed and delayed in Wallerian degeneration slow (Wld<jats:sup>S<\/jats:sup>) axons and this phenotype could be reproduced by intra-axonal disulfide reduction in wild-type axons and reversed by extra-axonal reduction in Wld<jats:sup>S<\/jats:sup>axons. These results are consistent with a redox mechanism for propagation of membrane polarity asymmetry in axonal degeneration.<\/jats:p>","DOI":"10.1523\/jneurosci.3882-16.2017","type":"journal-article","created":{"date-parts":[[2017,3,9]],"date-time":"2017-03-09T02:05:24Z","timestamp":1489025124000},"page":"3824-3839","source":"Crossref","is-referenced-by-count":31,"title":["Axonal Degeneration in Retinal Ganglion Cells Is Associated with a Membrane Polarity-Sensitive Redox Process"],"prefix":"10.1523","volume":"37","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1533-6642","authenticated-orcid":false,"given":"Mohammadali","family":"Almasieh","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria-Magdalena","family":"Catrinescu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lo\u00efc","family":"Binan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Santiago","family":"Costantino","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1399-250X","authenticated-orcid":false,"given":"Leonard A.","family":"Levin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"393","published-online":{"date-parts":[[2017,3,8]]},"reference":[{"key":"2023041803243094000_37.14.3824.1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.2004.03833.x"},{"key":"2023041803243094000_37.14.3824.2","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroscience.2012.08.056"},{"key":"2023041803243094000_37.14.3824.3","doi-asserted-by":"publisher","DOI":"10.1111\/j.1471-4159.2011.07382.x"},{"key":"2023041803243094000_37.14.3824.4","doi-asserted-by":"publisher","DOI":"10.1111\/j.1600-0854.2009.00974.x"},{"key":"2023041803243094000_37.14.3824.5","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2012.02.043"},{"key":"2023041803243094000_37.14.3824.6","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M700202200"},{"key":"2023041803243094000_37.14.3824.7","doi-asserted-by":"publisher","DOI":"10.1016\/0896-6273(88)90127-4"},{"key":"2023041803243094000_37.14.3824.8","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4065-10.2011"},{"key":"2023041803243094000_37.14.3824.9","doi-asserted-by":"publisher","DOI":"10.1038\/296150a0"},{"key":"2023041803243094000_37.14.3824.10","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1152\/jn.1993.69.1.151","article-title":"Alterations in membrane potential after axotomy at different distances from the soma of an identified neuron and the effect of depolarization on neurite outgrowth and calcium channel expression","volume":"69","author":"Berdan","year":"1993","journal-title":"J Neurophysiol"},{"key":"2023041803243094000_37.14.3824.11","doi-asserted-by":"publisher","DOI":"10.1016\/j.febslet.2010.03.020"},{"key":"2023041803243094000_37.14.3824.12","doi-asserted-by":"publisher","DOI":"10.1038\/nrn3710"},{"key":"2023041803243094000_37.14.3824.13","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.1995.tb01155.x"},{"key":"2023041803243094000_37.14.3824.14","doi-asserted-by":"publisher","DOI":"10.1016\/j.yexcr.2011.07.002"},{"key":"2023041803243094000_37.14.3824.15","doi-asserted-by":"publisher","DOI":"10.1016\/j.brainres.2004.01.036"},{"key":"2023041803243094000_37.14.3824.16","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-neuro-060909-153248"},{"key":"2023041803243094000_37.14.3824.17","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.95.17.9985"},{"key":"2023041803243094000_37.14.3824.18","doi-asserted-by":"publisher","DOI":"10.1021\/bi00453a005"},{"key":"2023041803243094000_37.14.3824.19","doi-asserted-by":"publisher","DOI":"10.1016\/0005-2736(91)90247-6"},{"key":"2023041803243094000_37.14.3824.20","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroscience.2011.05.052"},{"key":"2023041803243094000_37.14.3824.21","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2141.2006.06045.x"},{"key":"2023041803243094000_37.14.3824.22","doi-asserted-by":"publisher","DOI":"10.1080\/10409230903193307"},{"key":"2023041803243094000_37.14.3824.23","doi-asserted-by":"publisher","DOI":"10.1006\/bbrc.1999.1820"},{"key":"2023041803243094000_37.14.3824.24","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M003649200"},{"key":"2023041803243094000_37.14.3824.25","doi-asserted-by":"crossref","first-page":"2318","DOI":"10.1523\/JNEUROSCI.16-07-02318.1996","article-title":"An intrinsic time limit between genesis and death of individual neurons in the developing retinal ganglion cell layer","volume":"16","author":"Galli-Resta","year":"1996","journal-title":"J Neurosci"},{"key":"2023041803243094000_37.14.3824.26","doi-asserted-by":"crossref","first-page":"6445","DOI":"10.1523\/JNEUROSCI.15-10-06445.1995","article-title":"Axotomy-induced axonal degeneration is mediated by calcium influx through ion-specific channels","volume":"15","author":"George","year":"1995","journal-title":"J Neurosci"},{"key":"2023041803243094000_37.14.3824.27","doi-asserted-by":"publisher","DOI":"10.1038\/nrm1661"},{"key":"2023041803243094000_37.14.3824.28","doi-asserted-by":"publisher","DOI":"10.1016\/S0896-6273(02)00602-5"},{"key":"2023041803243094000_37.14.3824.29","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms6115"},{"key":"2023041803243094000_37.14.3824.30","doi-asserted-by":"publisher","DOI":"10.1167\/iovs.09-4450"},{"key":"2023041803243094000_37.14.3824.31","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M402568200"},{"key":"2023041803243094000_37.14.3824.32","doi-asserted-by":"publisher","DOI":"10.1021\/bi051751m"},{"key":"2023041803243094000_37.14.3824.33","doi-asserted-by":"publisher","DOI":"10.1016\/S0014-5793(00)01707-5"},{"key":"2023041803243094000_37.14.3824.34","doi-asserted-by":"publisher","DOI":"10.1021\/bi0107492"},{"key":"2023041803243094000_37.14.3824.35","doi-asserted-by":"publisher","DOI":"10.1093\/brain\/awq105"},{"key":"2023041803243094000_37.14.3824.36","doi-asserted-by":"publisher","DOI":"10.1038\/nm1229"},{"key":"2023041803243094000_37.14.3824.37","doi-asserted-by":"publisher","DOI":"10.1038\/nmeth.1405"},{"key":"2023041803243094000_37.14.3824.38","doi-asserted-by":"publisher","DOI":"10.1038\/nprot.2010.101"},{"key":"2023041803243094000_37.14.3824.39","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0909794107"},{"key":"2023041803243094000_37.14.3824.40","doi-asserted-by":"publisher","DOI":"10.1006\/exnr.1999.7014"},{"key":"2023041803243094000_37.14.3824.41","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.95.7.3966"},{"key":"2023041803243094000_37.14.3824.42","doi-asserted-by":"publisher","DOI":"10.1093\/emboj\/18.7.1761"},{"key":"2023041803243094000_37.14.3824.43","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.biophys.093008.131234"},{"key":"2023041803243094000_37.14.3824.44","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2016.07.064"},{"key":"2023041803243094000_37.14.3824.45","doi-asserted-by":"publisher","DOI":"10.1007\/s00441-012-1362-3"},{"key":"2023041803243094000_37.14.3824.46","doi-asserted-by":"publisher","DOI":"10.1111\/j.1471-4159.1990.tb13317.x"},{"key":"2023041803243094000_37.14.3824.47","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.1989.tb00771.x"},{"key":"2023041803243094000_37.14.3824.48","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.neuro.28.061604.135632"},{"key":"2023041803243094000_37.14.3824.49","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.90.20.9717"},{"key":"2023041803243094000_37.14.3824.50","doi-asserted-by":"publisher","DOI":"10.1038\/nn770"},{"key":"2023041803243094000_37.14.3824.51","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1097-4547(20000215)59:4<568::AID-JNR13>3.0.CO;2-R"},{"key":"2023041803243094000_37.14.3824.52","doi-asserted-by":"publisher","DOI":"10.1084\/jem.182.5.1545"},{"key":"2023041803243094000_37.14.3824.53","first-page":"119","article-title":"Oxidized phosphatidylserine: production and bioactivities","volume":"57","author":"Matsura","year":"2014","journal-title":"Yonago Acta Med"},{"key":"2023041803243094000_37.14.3824.54","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1013965108"},{"key":"2023041803243094000_37.14.3824.55","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuint.2016.05.014"},{"key":"2023041803243094000_37.14.3824.56","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1331-13.2013"},{"key":"2023041803243094000_37.14.3824.57","doi-asserted-by":"publisher","DOI":"10.1080\/03602530701468516"},{"key":"2023041803243094000_37.14.3824.58","doi-asserted-by":"publisher","DOI":"10.1002\/brb3.33"},{"key":"2023041803243094000_37.14.3824.59","doi-asserted-by":"publisher","DOI":"10.1021\/bi047642+"},{"key":"2023041803243094000_37.14.3824.60","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.1990.tb00472.x"},{"key":"2023041803243094000_37.14.3824.61","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.1991.tb00815.x"},{"key":"2023041803243094000_37.14.3824.62","doi-asserted-by":"publisher","DOI":"10.1002\/jnr.10176"},{"key":"2023041803243094000_37.14.3824.63","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1242\/dev.112.1.83","article-title":"The effect of age on motor neurone death following axotomy in the mouse","volume":"112","author":"Pollin","year":"1991","journal-title":"Development"},{"key":"2023041803243094000_37.14.3824.64","doi-asserted-by":"publisher","DOI":"10.1007\/s00018-006-6167-7"},{"key":"2023041803243094000_37.14.3824.65","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1016\/0165-3806(82)90013-X","article-title":"The loss of ganglion cells in the developing retina of the rat","volume":"255","author":"Potts","year":"1982","journal-title":"Brain Res"},{"key":"2023041803243094000_37.14.3824.66","doi-asserted-by":"publisher","DOI":"10.1126\/science.1068613"},{"key":"2023041803243094000_37.14.3824.67","doi-asserted-by":"publisher","DOI":"10.1096\/fj.05-5460rev"},{"key":"2023041803243094000_37.14.3824.68","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1097-4547(19970615)48:6<563::AID-JNR9>3.0.CO;2-8"},{"key":"2023041803243094000_37.14.3824.69","doi-asserted-by":"publisher","DOI":"10.1038\/nrn3609"},{"key":"2023041803243094000_37.14.3824.70","doi-asserted-by":"publisher","DOI":"10.1126\/science.402030"},{"key":"2023041803243094000_37.14.3824.71","doi-asserted-by":"publisher","DOI":"10.1016\/j.exer.2006.07.002"},{"key":"2023041803243094000_37.14.3824.72","doi-asserted-by":"publisher","DOI":"10.1126\/science.1252809"},{"key":"2023041803243094000_37.14.3824.73","doi-asserted-by":"publisher","DOI":"10.1016\/S0168-0102(03)00039-7"},{"key":"2023041803243094000_37.14.3824.74","doi-asserted-by":"publisher","DOI":"10.1155\/2016\/1087647"},{"key":"2023041803243094000_37.14.3824.75","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M611090200"},{"key":"2023041803243094000_37.14.3824.76","doi-asserted-by":"publisher","DOI":"10.1038\/nature09583"},{"key":"2023041803243094000_37.14.3824.77","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.bb.23.060194.001205"},{"key":"2023041803243094000_37.14.3824.78","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1509879112"},{"key":"2023041803243094000_37.14.3824.79","doi-asserted-by":"publisher","DOI":"10.1385\/CBB:38:2:161"},{"key":"2023041803243094000_37.14.3824.80","doi-asserted-by":"crossref","first-page":"1783","DOI":"10.1242\/jcs.107.7.1783","article-title":"The mode of anchorage to the cell surface determines both the function and the membrane location of Thy-1 glycoprotein","volume":"107","author":"Tiveron","year":"1994","journal-title":"J Cell Sci"},{"key":"2023041803243094000_37.14.3824.81","doi-asserted-by":"crossref","first-page":"4718","DOI":"10.1523\/JNEUROSCI.19-12-04718.1999","article-title":"Temperature modulation reveals three distinct stages of Wallerian degeneration","volume":"19","author":"Tsao","year":"1999","journal-title":"J Neurosci"},{"key":"2023041803243094000_37.14.3824.82","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbrc.2004.09.102"},{"key":"2023041803243094000_37.14.3824.83","doi-asserted-by":"publisher","DOI":"10.1038\/nrm2330"},{"key":"2023041803243094000_37.14.3824.84","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2484-15.2015"},{"key":"2023041803243094000_37.14.3824.85","doi-asserted-by":"publisher","DOI":"10.1016\/j.preteyeres.2015.06.003"},{"key":"2023041803243094000_37.14.3824.86","doi-asserted-by":"publisher","DOI":"10.1016\/0005--2736(73)90143--0"},{"key":"2023041803243094000_37.14.3824.87","doi-asserted-by":"publisher","DOI":"10.1002\/neu.480240103"},{"key":"2023041803243094000_37.14.3824.88","doi-asserted-by":"publisher","DOI":"10.1098\/rstl.1850.0021"},{"key":"2023041803243094000_37.14.3824.89","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.201108111"},{"key":"2023041803243094000_37.14.3824.90","doi-asserted-by":"publisher","DOI":"10.4137\/LPI.S31785"},{"key":"2023041803243094000_37.14.3824.91","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1242\/dev.112.1.161","article-title":"The surface glycoprotein Thy-1 is excluded from growing axons during development: a study of the expression of Thy-1 during axogenesis in hippocampus and hindbrain","volume":"112","author":"Xue","year":"1991","journal-title":"Development"},{"key":"2023041803243094000_37.14.3824.92","doi-asserted-by":"publisher","DOI":"10.1089\/ars.2007.1672"},{"key":"2023041803243094000_37.14.3824.93","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.1993.tb00531.x"}],"container-title":["The Journal of Neuroscience"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1523\/JNEUROSCI.3882-16.2017","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,4,18]],"date-time":"2023-04-18T10:57:34Z","timestamp":1681815454000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.jneurosci.org\/lookup\/doi\/10.1523\/JNEUROSCI.3882-16.2017"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2017,3,8]]},"references-count":93,"journal-issue":{"issue":"14","published-online":{"date-parts":[[2017,4,5]]},"published-print":{"date-parts":[[2017,4,5]]}},"alternative-id":["10.1523\/JNEUROSCI.3882-16.2017"],"URL":"https:\/\/doi.org\/10.1523\/jneurosci.3882-16.2017","relation":{},"ISSN":["0270-6474","1529-2401"],"issn-type":[{"value":"0270-6474","type":"print"},{"value":"1529-2401","type":"electronic"}],"subject":[],"published":{"date-parts":[[2017,3,8]]}}}