{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,29]],"date-time":"2026-09-29T09:36:35Z","timestamp":1790674595210,"version":"4.1.0"},"reference-count":70,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2020,4,2]],"date-time":"2020-04-02T00:00:00Z","timestamp":1585785600000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2020,4,2]],"date-time":"2020-04-02T00:00:00Z","timestamp":1585785600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"CHU de Quebec-Universite Laval","award":["start-up fund"],"award-info":[{"award-number":["start-up fund"]}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["J Neuroinflammation"],"published-print":{"date-parts":[[2020,12]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>\n                      Huntington\u2019s disease (HD) is a dominantly inherited neurodegenerative disorder that affects cognitive and motor abilities by primarily targeting the striatum and cerebral cortex. HD is caused by a mutation elongating the CAG repeats within the\n                      <jats:italic>Huntingtin<\/jats:italic>\n                      gene, resulting in HTT protein misfolding. Although the genetic cause of HD has been established, the specific susceptibility of neurons within various brain structures has remained elusive. Microglia, which are the brain\u2019s resident macrophages, have emerged as important players in neurodegeneration. Nevertheless, few studies have examined their implication in HD.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>To provide novel insights, we investigated the maturation and dysfunction of striatal microglia using the R6\/2 mouse model of HD. This transgenic model, which presents with 120+\/-5 CAG repeats, displays progressive motor deficits beginning at 6\u2009weeks of age, with full incapacitation by 13\u2009weeks. We studied microglial morphology, phagocytic capacity, and synaptic contacts in the striatum of R6\/2 versus wild-type (WT) littermates at 3, 10, and 13\u2009weeks of age, using a combination of light and transmission electron microscopy. We also reconstructed dendrites and determined synaptic density within the striatum of R6\/2 and WT littermates, at nanoscale resolution using focused ion beam scanning electron microscopy.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>At 3\u2009weeks of age, prior to any known motor deficits, microglia in R6\/2 animals displayed a more mature morphological phenotype than WT animals. Microglia from R6\/2 mice across all ages also demonstrated increased phagocytosis, as revealed by light microscopy and transmission electron microscopy. Furthermore, microglial processes from 10-week-old R6\/2 mice made fewer contacts with synaptic structures than microglial processes in 3-week-old R6\/2 mice and age-matched WT littermates. Synaptic density was not affected by genotype at 3\u2009weeks of age but increased with maturation in WT mice. The location of synapses was lastly modified in R6\/2 mice compared with WT controls, from targeting dendritic spines to dendritic trunks at both 3 and 10\u2009weeks of age.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>These findings suggest that microglia may play an intimate role in synaptic alteration and loss during HD pathogenesis.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/s12974-020-01782-9","type":"journal-article","created":{"date-parts":[[2020,4,2]],"date-time":"2020-04-02T08:02:40Z","timestamp":1585814560000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":102,"title":["Microglial physiological properties and interactions with synapses are altered at presymptomatic stages in a mouse model of Huntington\u2019s disease pathology"],"prefix":"10.1186","volume":"17","author":[{"given":"Julie C.","family":"Savage","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marie-Kim","family":"St-Pierre","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mica\u00ebl","family":"Carrier","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hassan","family":"El Hajj","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sammy Weiser","family":"Novak","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria Gabriela","family":"Sanchez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Francesca","family":"Cicchetti","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marie-\u00c8ve","family":"Tremblay","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2020,4,2]]},"reference":[{"key":"1782_CR1","doi-asserted-by":"publisher","first-page":"204","DOI":"10.1038\/nrneurol.2014.24","volume":"10","author":"CA Ross","year":"2014","unstructured":"Ross CA, Aylward EH, Wild EJ, Langbehn DR, Long JD, Warner JH, et al. Huntington disease: natural history, biomarkers and prospects for therapeutics. Nat Rev Neurol. 2014;10:204\u201316.","journal-title":"Nat Rev Neurol"},{"key":"1782_CR2","doi-asserted-by":"publisher","first-page":"971","DOI":"10.1016\/0092-8674(93)90585-E","volume":"72","author":"ME MacDonald","year":"1993","unstructured":"MacDonald ME, Ambrose CM, Duyao MP, Myers RH, Lin C, Srinidhi L, et al. A novel gene containing a trinucleotide repeat that is expanded and unstable on Huntington\u2019s disease chromosomes. Cell. 1993;72:971\u201383.","journal-title":"Cell"},{"key":"1782_CR3","doi-asserted-by":"publisher","first-page":"407","DOI":"10.1126\/science.7618107","volume":"269","author":"MP Duyao","year":"1995","unstructured":"Duyao MP, Auerbach AB, Ryan A, Persichetti F, Barnes GT, McNeil SM, et al. Inactivation of the mouse Huntington\u2019s disease gene homolog Hdh. Science. 1995;269:407\u201310.","journal-title":"Science"},{"key":"1782_CR4","doi-asserted-by":"publisher","first-page":"811","DOI":"10.1016\/0092-8674(95)90542-1","volume":"81","author":"J Nasir","year":"1995","unstructured":"Nasir J, Floresco SB, O\u2019Kusky JR, Diewert VM, Richman JM, Zeisler J, et al. Targeted disruption of the Huntington\u2019s disease gene results in embryonic lethality and behavioral and morphological changes in heterozygotes. Cell. 1995;81:811\u201323.","journal-title":"Cell"},{"key":"1782_CR5","first-page":"16","volume":"59","author":"DC Rubinsztein","year":"1996","unstructured":"Rubinsztein DC, Leggo J, Coles R, Almqvist E, Biancalana V, Cassiman JJ, et al. Phenotypic characterization of individuals with 30-40 CAG repeats in the Huntington disease (HD) gene reveals HD cases with 36 repeats and apparently normal elderly individuals with 36-39 repeats. Am J Hum Genet. 1996;59:16\u201322.","journal-title":"Am J Hum Genet"},{"key":"1782_CR6","doi-asserted-by":"publisher","first-page":"1990","DOI":"10.1126\/science.277.5334.1990","volume":"277","author":"M DiFiglia","year":"1997","unstructured":"DiFiglia M, Sapp E, Chase KO, Davies SW, Bates GP, Vonsattel JP, et al. Aggregation of Huntingtin in neuronal intranuclear inclusions and dystrophic neurites in brain. Science. 1997;277:1990\u20133.","journal-title":"Science"},{"key":"1782_CR7","doi-asserted-by":"publisher","first-page":"398","DOI":"10.1038\/ng0893-398","volume":"4","author":"SE Andrew","year":"1993","unstructured":"Andrew SE, Goldberg YP, Kremer B, Telenius H, Theilmann J, Adam S, et al. The relationship between trinucleotide (CAG) repeat length and clinical features of Huntington\u2019s disease. Nat Genet. 1993;4:398\u2013403.","journal-title":"Nat Genet"},{"key":"1782_CR8","doi-asserted-by":"publisher","first-page":"493","DOI":"10.1016\/S0092-8674(00)81369-0","volume":"87","author":"L Mangiarini","year":"1996","unstructured":"Mangiarini L, Sathasivam K, Seller M, Cozens B, Harper A, Hetherington C, et al. Exon 1 of the HD gene with an expanded CAG repeat is sufficient to cause a progressive neurological phenotype in transgenic mice. Cell. 1996;87:493\u2013506.","journal-title":"Cell"},{"key":"1782_CR9","doi-asserted-by":"publisher","first-page":"5733","DOI":"10.1073\/pnas.85.15.5733","volume":"85","author":"A Reiner","year":"1988","unstructured":"Reiner A, Albin RL, Anderson KD, D\u2019Amato CJ, Penney JB, Young AB. Differential loss of striatal projection neurons in Huntington disease. Proc Natl Acad Sci U S A. 1988;85:5733\u20137.","journal-title":"Proc Natl Acad Sci U S A"},{"key":"1782_CR10","doi-asserted-by":"publisher","first-page":"270","DOI":"10.1007\/s13311-012-0112-2","volume":"9","author":"ME Ehrlich","year":"2012","unstructured":"Ehrlich ME. Huntington\u2019s disease and the striatal medium spiny neuron: cell-autonomous and non-cell-autonomous mechanisms of disease. Neurotherapeutics. 2012;9:270\u201384.","journal-title":"Neurotherapeutics"},{"key":"1782_CR11","doi-asserted-by":"publisher","first-page":"1615","DOI":"10.1212\/01.WNL.0000065888.88988.6E","volume":"60","author":"HD Rosas","year":"2003","unstructured":"Rosas HD, Koroshetz WJ, Chen YI, Skeuse C, Vangel M, Cudkowicz ME, et al. Evidence for more widespread cerebral pathology in early HD: an MRI-based morphometric analysis. Neurology. 2003;60:1615\u201320.","journal-title":"Neurology"},{"key":"1782_CR12","doi-asserted-by":"publisher","first-page":"559","DOI":"10.1097\/00005072-198511000-00003","volume":"44","author":"JP Vonsattel","year":"1985","unstructured":"Vonsattel JP, Myers RH, Stevens TJ, Ferrante RJ, Bird ED, Richardson EP. Neuropathological classification of Huntington\u2019s disease. J Neuropathol Exp Neurol. 1985;44:559\u201377.","journal-title":"J Neuropathol Exp Neurol"},{"key":"1782_CR13","doi-asserted-by":"publisher","first-page":"8093","DOI":"10.1073\/pnas.110078997","volume":"97","author":"M Turmaine","year":"2000","unstructured":"Turmaine M, Raza A, Mahal A, Mangiarini L, Bates GP, Davies SW. Nonapoptotic neurodegeneration in a transgenic mouse model of Huntington\u2019s disease. Proc Natl Acad Sci. 2000;97:8093\u20137.","journal-title":"Proc Natl Acad Sci"},{"key":"1782_CR14","doi-asserted-by":"publisher","first-page":"961","DOI":"10.1523\/JNEUROSCI.23-03-00961.2003","volume":"23","author":"C Cepeda","year":"2003","unstructured":"Cepeda C, Hurst RS, Calvert CR, Hern\u00e1ndez-Echeagaray E, Nguyen OK, Jocoy E, et al. Transient and progressive electrophysiological alterations in the corticostriatal pathway in a mouse model of Huntington\u2019s disease. J Neurosci. 2003;23:961\u20139.","journal-title":"J Neurosci"},{"key":"1782_CR15","doi-asserted-by":"publisher","first-page":"863","DOI":"10.1016\/j.cell.2011.05.020","volume":"145","author":"D Zwilling","year":"2011","unstructured":"Zwilling D, Huang S-Y, Sathyasaikumar KV, Notarangelo FM, Guidetti P, Wu H-Q, et al. Kynurenine 3-monooxygenase inhibition in blood ameliorates neurodegeneration. Cell. 2011;145:863\u201374.","journal-title":"Cell"},{"key":"1782_CR16","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1523\/JNEUROSCI.0244-14.2015","volume":"35","author":"RP Murmu","year":"2015","unstructured":"Murmu RP, Li W, Szepesi Z, Li J-Y. Altered sensory experience exacerbates stable dendritic spine and synapse loss in a mouse model of Huntington\u2019s disease. J Neurosci. 2015;35:287\u201398.","journal-title":"J Neurosci"},{"key":"1782_CR17","doi-asserted-by":"publisher","first-page":"50","DOI":"10.1002\/glia.23050","volume":"65","author":"AHP Jansen","year":"2017","unstructured":"Jansen AHP, Van Hal M, Op den Kelder IC, Meier RT, de Ruiter A-A, Schut MH, et al. Frequency of nuclear mutant huntingtin inclusion formation in neurons and glia is cell-type-specific. Glia. 2017;65:50\u201361.","journal-title":"Glia"},{"key":"1782_CR18","doi-asserted-by":"publisher","first-page":"22480","DOI":"10.1073\/pnas.0911503106","volume":"106","author":"J Bradford","year":"2009","unstructured":"Bradford J, Shin J-Y, Roberts M, Wang C-E, Li X-J, Li S. Expression of mutant huntingtin in mouse brain astrocytes causes age-dependent neurological symptoms. Proc Natl Acad Sci U S A. 2009;106:22480\u20135.","journal-title":"Proc Natl Acad Sci U S A."},{"key":"1782_CR19","doi-asserted-by":"publisher","first-page":"1212","DOI":"10.1016\/j.neuron.2015.02.026","volume":"85","author":"B Huang","year":"2015","unstructured":"Huang B, Wei W, Wang G, Gaertig MA, Feng Y, Wang W, et al. Mutant huntingtin downregulates myelin regulatory factor-mediated myelin gene expression and affects mature oligodendrocytes. Neuron. 2015;85:1212\u201326.","journal-title":"Neuron"},{"key":"1782_CR20","doi-asserted-by":"publisher","first-page":"128","DOI":"10.1016\/j.conb.2015.12.004","volume":"36","author":"S Hong","year":"2016","unstructured":"Hong S, Dissing-Olesen L, Stevens B. New insights on the role of microglia in synaptic pruning in health and disease. Curr Opin Neurobiol. 2016;36:128\u201334.","journal-title":"Curr Opin Neurobiol"},{"key":"1782_CR21","doi-asserted-by":"crossref","unstructured":"Henstridge CM, Tzioras M, Paolicelli RC. Glial contribution to excitatory and inhibitory synapse loss in neurodegeneration. Front Cell Neurosci. 2019 [cited 2019 Dec 12];13. Available from: https:\/\/www.frontiersin.org\/articles\/10.3389\/fncel.2019.00063\/full.","DOI":"10.3389\/fncel.2019.00063"},{"key":"1782_CR22","doi-asserted-by":"crossref","unstructured":"Yang H-M, Yang S, Huang S-S, Tang B-S, Guo J-F. Microglial activation in the pathogenesis of Huntington\u2019s disease. Front Aging Neurosci. 2017 [cited 2019 Oct 22];9. Available from: https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC5474461\/.","DOI":"10.3389\/fnagi.2017.00193"},{"key":"1782_CR23","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1093\/jnen\/60.2.161","volume":"60","author":"E Sapp","year":"2001","unstructured":"Sapp E, Kegel KB, Aronin N, Hashikawa T, Uchiyama Y, Tohyama K, et al. Early and progressive accumulation of reactive microglia in the Huntington disease brain. J Neuropathol Exp Neurol. 2001;60:161\u201372.","journal-title":"J Neuropathol Exp Neurol."},{"key":"1782_CR24","doi-asserted-by":"publisher","first-page":"438","DOI":"10.1016\/j.nbd.2011.09.003","volume":"45","author":"S Franciosi","year":"2012","unstructured":"Franciosi S, Ryu JK, Shim Y, Hill A, Connolly C, Hayden MR, et al. Age-dependent neurovascular abnormalities and altered microglial morphology in the YAC128 mouse model of Huntington disease. Neurobiol Dis. 2012;45:438\u201349.","journal-title":"Neurobiol Dis"},{"key":"1782_CR25","doi-asserted-by":"publisher","first-page":"621.e17","DOI":"10.1016\/j.neurobiolaging.2011.02.015","volume":"33","author":"AD Kraft","year":"2012","unstructured":"Kraft AD, Kaltenbach LS, Lo DC, Harry GJ. Activated microglia proliferate at neurites of mutant huntingtin-expressing neurons. Neurobiol Aging. 2012;33:621.e17\u201333.","journal-title":"Neurobiol Aging"},{"key":"1782_CR26","doi-asserted-by":"publisher","first-page":"1869","DOI":"10.1084\/jem.20080178","volume":"205","author":"M Bj\u00f6rkqvist","year":"2008","unstructured":"Bj\u00f6rkqvist M, Wild EJ, Thiele J, Silvestroni A, Andre R, Lahiri N, et al. A novel pathogenic pathway of immune activation detectable before clinical onset in Huntington\u2019s disease. J Exp Med. 2008;205:1869\u201377.","journal-title":"J Exp Med"},{"key":"1782_CR27","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1038\/nn.3668","volume":"17","author":"A Crotti","year":"2014","unstructured":"Crotti A, Benner C, Kerman BE, Gosselin D, Lagier-Tourenne C, Zuccato C, et al. Mutant Huntingtin promotes autonomous microglia activation via myeloid lineage-determining factors. Nat Neurosci. 2014;17:513\u201321.","journal-title":"Nat Neurosci"},{"key":"1782_CR28","doi-asserted-by":"publisher","first-page":"1638","DOI":"10.1212\/01.wnl.0000222734.56412.17","volume":"66","author":"N Pavese","year":"2006","unstructured":"Pavese N, Gerhard A, Tai YF, Ho AK, Turkheimer F, Barker RA, et al. Microglial activation correlates with severity in Huntington disease: a clinical and PET study. Neurology. 2006;66:1638\u201343.","journal-title":"Neurology"},{"key":"1782_CR29","doi-asserted-by":"publisher","first-page":"1759","DOI":"10.1093\/brain\/awm044","volume":"130","author":"YF Tai","year":"2007","unstructured":"Tai YF, Pavese N, Gerhard A, Tabrizi SJ, Barker RA, Brooks DJ, et al. Microglial activation in presymptomatic Huntington\u2019s disease gene carriers. Brain. 2007;130:1759\u201366.","journal-title":"Brain"},{"key":"1782_CR30","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1016\/j.nbd.2015.08.011","volume":"83","author":"M Politis","year":"2015","unstructured":"Politis M, Lahiri N, Niccolini F, Su P, Wu K, Giannetti P, et al. Increased central microglial activation associated with peripheral cytokine levels in premanifest Huntington\u2019s disease gene carriers. Neurobiol Dis. 2015;83:115\u201321.","journal-title":"Neurobiol Dis"},{"key":"1782_CR31","doi-asserted-by":"publisher","first-page":"461","DOI":"10.1016\/S0140-6736(01)05625-2","volume":"358","author":"A Cagnin","year":"2001","unstructured":"Cagnin A, Brooks DJ, Kennedy AM, Gunn RN, Myers R, Turkheimer FE, et al. In-vivo measurement of activated microglia in dementia. Lancet. 2001;358:461\u20137.","journal-title":"Lancet"},{"key":"1782_CR32","doi-asserted-by":"publisher","first-page":"362","DOI":"10.1006\/exnr.1999.7170","volume":"159","author":"SK Singhrao","year":"1999","unstructured":"Singhrao SK, Neal JW, Morgan BP, Gasque P. Increased complement biosynthesis by microglia and complement activation on neurons in Huntington\u2019s disease. Exp Neurol. 1999;159:362\u201376.","journal-title":"Exp Neurol"},{"key":"1782_CR33","doi-asserted-by":"publisher","first-page":"1929","DOI":"10.1113\/JP272134","volume":"595","author":"TL Tay","year":"2017","unstructured":"Tay TL, Savage JC, Hui CW, Bisht K, Tremblay M-\u00c8. Microglia across the lifespan: from origin to function in brain development, plasticity and cognition. J Physiol. 2017;595:1929\u201345.","journal-title":"J Physiol"},{"key":"1782_CR34","doi-asserted-by":"publisher","first-page":"250","DOI":"10.1111\/cns.12844","volume":"24","author":"A Reiner","year":"2018","unstructured":"Reiner A, Deng Y-P. Disrupted striatal neuron inputs and outputs in Huntington\u2019s disease. CNS Neurosci Ther. 2018;24:250\u201380.","journal-title":"CNS Neurosci Ther"},{"key":"1782_CR35","doi-asserted-by":"publisher","first-page":"e75099","DOI":"10.1371\/journal.pone.0075099","volume":"8","author":"S Bissonnette","year":"2013","unstructured":"Bissonnette S, Vaillancourt M, H\u00e9bert SS, Drolet G, Samadi P. Striatal pre-enkephalin overexpression improves Huntington\u2019s disease symptoms in the R6\/2 mouse model of Huntington\u2019s disease. Plos One. 2013;8:e75099.","journal-title":"Plos One"},{"key":"1782_CR36","doi-asserted-by":"crossref","unstructured":"Bisht K, El Hajj H, Savage JC, S\u00e1nchez MG, Tremblay M-\u00c8. Correlative light and electron microscopy to study microglial interactions with \u03b2-amyloid plaques. J Vis Exp JoVE. 2016.","DOI":"10.3791\/54060"},{"key":"1782_CR37","doi-asserted-by":"publisher","first-page":"e1000527","DOI":"10.1371\/journal.pbio.1000527","volume":"8","author":"M-\u00c8 Tremblay","year":"2010","unstructured":"Tremblay M-\u00c8, Lowery RL, Majewska AK. Microglial interactions with synapses are modulated by visual experience. PLoS Biol. 2010;8:e1000527.","journal-title":"PLoS Biol"},{"key":"1782_CR38","unstructured":"Deerinck TJ, Bushong EA, Thor A, Ellisman MH. NCMIR Methods for 3D EM: a new protocol for preparation of biological specimens for serial blockface scanning electron microscopy. [cited 2019 Nov 15]. Available from: https:\/\/ncmir.ucsd.edu\/sbem-protocol\/."},{"key":"1782_CR39","doi-asserted-by":"publisher","first-page":"450","DOI":"10.1016\/j.bbi.2018.06.007","volume":"73","author":"CW Hui","year":"2018","unstructured":"Hui CW, St-Pierre M-K, Detuncq J, Aumailley L, Dubois M-J, Couture V, et al. Nonfunctional mutant Wrn protein leads to neurological deficits, neuronal stress, microglial alteration, and immune imbalance in a mouse model of Werner syndrome. Brain Behav Immun. 2018;73:450\u201369.","journal-title":"Brain Behav Immun"},{"key":"1782_CR40","unstructured":"Ibanez FG, Picard K, Bordelau M, Sharma K, Bisht K, Tremblay M-\u00c8. Immunofluorescence staining using IBA1 and TMEM119 for microglial density, morphology and peripheral myeloid cell infiltration analysis in mouse brain. JoVE J Vis Exp. 2019:e60510."},{"key":"1782_CR41","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","volume":"9","author":"J Schindelin","year":"2012","unstructured":"Schindelin J, Arganda-Carreras I, Frise E, Kaynig V, Longair M, Pietzsch T, et al. Fiji: an open-source platform for biological-image analysis. Nat Methods. 2012;9:676\u201382.","journal-title":"Nat Methods"},{"key":"1782_CR42","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1002\/glia.22287","volume":"60","author":"M-\u00c8 Tremblay","year":"2012","unstructured":"Tremblay M-\u00c8, Zettel ML, Ison JR, Allen PD, Majewska AK. Effects of aging and sensory loss on glial cells in mouse visual and auditory cortices. Glia. 2012;60:541\u201358.","journal-title":"Glia"},{"key":"1782_CR43","unstructured":"Peters A, Palay SL, Webster HD. Fine structures of the nervous system: neurons and their supporting cells. 3rd ed. Oxford University Press; 1991 [cited 2019 Nov 23]. Available from: https:\/\/www.amazon.com\/Fine-Structure-Nervous-System-Supporting\/dp\/0195065719."},{"key":"1782_CR44","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1186\/s12974-019-1473-9","volume":"16","author":"H El Hajj","year":"2019","unstructured":"El Hajj H, Savage JC, Bisht K, Parent M, Valli\u00e8res L, Rivest S, et al. Ultrastructural evidence of microglial heterogeneity in Alzheimer\u2019s disease amyloid pathology. J Neuroinflammation. 2019;16:87.","journal-title":"J Neuroinflammation"},{"key":"1782_CR45","doi-asserted-by":"crossref","unstructured":"Knott G, Rosset S, Cantoni M. Focussed ion beam milling and scanning electron microscopy of brain tissue. J Vis Exp JoVE [Internet]. 2011 [cited 2019 Nov 15]; Available from: https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC3196160\/.","DOI":"10.3791\/2588"},{"key":"1782_CR46","doi-asserted-by":"publisher","first-page":"1226","DOI":"10.1038\/s41592-019-0582-9","volume":"16","author":"S Berg","year":"2019","unstructured":"Berg S, Kutra D, Kroeger T, Straehle CN, Kausler BX, Haubold C, et al. ilastik: interactive machine learning for (bio)image analysis. Nat Methods. 2019;16:1226\u201332.","journal-title":"Nat Methods"},{"key":"1782_CR47","doi-asserted-by":"crossref","unstructured":"Merch\u00e1n-P\u00e9rez A, Rodriguez J-R, Alonso-Nanclares L, Schertel A, De Felipe J. Counting synapses using FIB\/SEM microscopy: a true revolution for ultrastructural volume reconstruction. Front Neuroanat. 2009 [cited 2020 Feb 7];3. Available from: https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC2784681\/.","DOI":"10.3389\/neuro.05.018.2009"},{"key":"1782_CR48","doi-asserted-by":"publisher","first-page":"e343","DOI":"10.1371\/journal.pbio.0040343","volume":"4","author":"C Genoud","year":"2006","unstructured":"Genoud C, Quairiaux C, Steiner P, Hirling H, Welker E, Knott GW. Plasticity of astrocytic coverage and glutamate transporter expression in adult mouse cortex. PLOS Biol. 2006;4:e343.","journal-title":"PLOS Biol"},{"key":"1782_CR49","doi-asserted-by":"crossref","unstructured":"Berger DR, Seung HS, Lichtman JW. VAST (Volume Annotation and Segmentation Tool): efficient manual and semi-automatic labeling of large 3D image stacks. Front Neural Circuits. 2018 [cited 2020 Feb 28];12. Available from: https:\/\/www.frontiersin.org\/articles\/10.3389\/fncir.2018.00088\/full.","DOI":"10.3389\/fncir.2018.00088"},{"key":"1782_CR50","doi-asserted-by":"publisher","first-page":"274","DOI":"10.1186\/1471-2105-11-274","volume":"11","author":"B Schmid","year":"2010","unstructured":"Schmid B, Schindelin J, Cardona A, Longair M, Heisenberg M. A high-level 3D visualization API for Java and ImageJ. BMC Bioinformatics. 2010;11:274.","journal-title":"BMC Bioinformatics"},{"key":"1782_CR51","volume-title":"Blender - a 3D modelling and rendering package","author":"BO Community","year":"2018","unstructured":"Community BO. Blender - a 3D modelling and rendering package. Amsterdam: Sttiching Blender Foundation; 2018. Available from: www.blender.org."},{"key":"1782_CR52","doi-asserted-by":"crossref","unstructured":"Jorstad A, Blanc J, Knott G. NeuroMorph: a software toolset for 3D analysis of neurite morphology and connectivity. Front Neuroanat. 2018 [cited 2020 Feb 28];12. Available from: https:\/\/www.ncbi.nlm.nih.gov\/pmc\/articles\/PMC6064741\/.","DOI":"10.3389\/fnana.2018.00059"},{"key":"1782_CR53","doi-asserted-by":"publisher","first-page":"354","DOI":"10.1002\/cne.20680","volume":"490","author":"EC Stack","year":"2005","unstructured":"Stack EC, Kubilus JK, Smith K, Cormier K, Signore SJD, Guelin E, et al. Chronology of behavioral symptoms and neuropathological sequela in R6\/2 Huntington\u2019s disease transgenic mice. J Comp Neurol. 2005;490:354\u201370.","journal-title":"J Comp Neurol"},{"key":"1782_CR54","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1016\/j.neulet.2019.01.001","volume":"698","author":"JO Uweru","year":"2019","unstructured":"Uweru JO, Eyo UB. A decade of diverse microglial-neuronal physical interactions in the brain (2008\u20132018). Neurosci Lett. 2019;698:33\u20138.","journal-title":"Neurosci Lett"},{"key":"1782_CR55","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1038\/labinvest.2016.116","volume":"97","author":"DA Chistiakov","year":"2017","unstructured":"Chistiakov DA, Killingsworth MC, Myasoedova VA, Orekhov AN, Bobryshev YV. CD68\/macrosialin: not just a histochemical marker. Lab Invest. 2017;97:4\u201313.","journal-title":"Lab Invest"},{"key":"1782_CR56","doi-asserted-by":"publisher","first-page":"346","DOI":"10.1097\/NEN.0b013e3181d44992","volume":"69","author":"S Sasaki","year":"2010","unstructured":"Sasaki S. Endoplasmic reticulum stress in motor neurons of the spinal cord in sporadic amyotrophic lateral sclerosis. J Neuropathol Exp Neurol. 2010;69:346\u201355.","journal-title":"J Neuropathol Exp Neurol"},{"key":"1782_CR57","doi-asserted-by":"publisher","first-page":"826","DOI":"10.1002\/glia.22966","volume":"64","author":"K Bisht","year":"2016","unstructured":"Bisht K, Sharma KP, Lecours C, S\u00e1nchez MG, El Hajj H, Milior G, et al. Dark microglia: a new phenotype predominantly associated with pathological states. Glia. 2016;64:826\u201339.","journal-title":"Glia"},{"key":"1782_CR58","doi-asserted-by":"crossref","unstructured":"Estrada-S\u00e1nchez AM, Rebec GV. Role of cerebral cortex in the neuropathology of Huntington\u2019s disease. Front Neural Circuits. 2013 [cited 2019 Nov 17];7. Available from: https:\/\/www.frontiersin.org\/articles\/10.3389\/fncir.2013.00019\/full#B140.","DOI":"10.3389\/fncir.2013.00019"},{"key":"1782_CR59","doi-asserted-by":"crossref","unstructured":"Nikodemova M, Kimyon RS, De I, Small AL, Collier LS, Watters JJ. Microglial numbers attain adult levels after undergoing a rapid decrease in cell number in the third postnatal week. J Neuroimmunol. 2015:280\u20138.","DOI":"10.1016\/j.jneuroim.2014.11.018"},{"key":"1782_CR60","doi-asserted-by":"publisher","first-page":"274","DOI":"10.1002\/glia.10261","volume":"43","author":"L Ma","year":"2003","unstructured":"Ma L, Morton AJ, Nicholson LFB. Microglia density decreases with age in a mouse model of Huntington\u2019s disease. Glia. 2003;43:274\u201380.","journal-title":"Glia"},{"key":"1782_CR61","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1038\/s41593-018-0100-x","volume":"21","author":"B Ajami","year":"2018","unstructured":"Ajami B, Samusik N, Wieghofer P, Ho PP, Crotti A, Bjornson Z, et al. Single-cell mass cytometry reveals distinct populations of brain myeloid cells in mouse neuroinflammation and neurodegeneration models. Nat Neurosci. 2018;21:541\u201351.","journal-title":"Nat Neurosci"},{"key":"1782_CR62","doi-asserted-by":"publisher","first-page":"12997","DOI":"10.1523\/JNEUROSCI.5284-12.2013","volume":"33","author":"RP Murmu","year":"2013","unstructured":"Murmu RP, Li W, Holtmaat A, Li J-Y. Dendritic spine instability leads to progressive neocortical spine loss in a mouse model of Huntington\u2019s disease. J Neurosci. 2013;33:12997\u20133009.","journal-title":"J Neurosci"},{"key":"1782_CR63","doi-asserted-by":"crossref","unstructured":"Jackson J, Jambrina E, Li J, Marston H, Menzies F, Phillips K, et al. Targeting the synapse in Alzheimer\u2019s disease. Front Neurosci. 2019 [cited 2020 Feb 10];13. Available from: https:\/\/www.frontiersin.org\/articles\/10.3389\/fnins.2019.00735\/full.","DOI":"10.3389\/fnins.2019.00735"},{"key":"1782_CR64","doi-asserted-by":"publisher","first-page":"995","DOI":"10.3233\/JAD-122038","volume":"34","author":"L Blazquez-Llorca","year":"2013","unstructured":"Blazquez-Llorca L, Merch\u00e1n-P\u00e9rez \u00c1, Rodr\u00edguez J-R, Gasc\u00f3n J, DeFelipe J. FIB\/SEM Technology and Alzheimer\u2019s disease: three-dimensional analysis of human cortical synapses. J Alzheimers Dis. 2013;34:995\u20131013.","journal-title":"J Alzheimers Dis"},{"key":"1782_CR65","doi-asserted-by":"crossref","unstructured":"Dom\u00ednguez-\u00c1lvaro M, Montero-Crespo M, Blazquez-Llorca L, De Felipe J, Alonso-Nanclares L. 3D electron microscopy study of synaptic organization of the normal human transentorhinal cortex and its possible alterations in Alzheimer\u2019s disease. eNeuro. 2019 [cited 2020 Feb 10];6. Available from: https:\/\/www.eneuro.org\/content\/6\/4\/ENEURO.0140-19.2019.","DOI":"10.1523\/ENEURO.0140-19.2019"},{"key":"1782_CR66","doi-asserted-by":"publisher","first-page":"1074","DOI":"10.1002\/glia.20526","volume":"55","author":"DA Simmons","year":"2007","unstructured":"Simmons DA, Casale M, Alcon B, Pham N, Narayan N, Lynch G. Ferritin accumulation in dystrophic microglia is an early event in the development of Huntington\u2019s disease. Glia. 2007;55:1074\u201384.","journal-title":"Glia"},{"key":"1782_CR67","doi-asserted-by":"publisher","first-page":"1661","DOI":"10.1093\/hmg\/ddz009","volume":"28","author":"TL Petkau","year":"2019","unstructured":"Petkau TL, Hill A, Connolly C, Lu G, Wagner P, Kosior N, et al. Mutant huntingtin expression in microglia is neither required nor sufficient to cause the Huntington\u2019s disease-like phenotype in BACHD mice. Hum Mol Genet. 2019;28:1661\u201370.","journal-title":"Hum Mol Genet"},{"key":"1782_CR68","doi-asserted-by":"publisher","first-page":"1001","DOI":"10.1083\/jcb.200508072","volume":"171","author":"J-Y Shin","year":"2005","unstructured":"Shin J-Y, Fang Z-H, Yu Z-X, Wang C-E, Li S-H, Li X-J. Expression of mutant huntingtin in glial cells contributes to neuronal excitotoxicity. J Cell Biol. 2005;171:1001\u201312.","journal-title":"J Cell Biol"},{"key":"1782_CR69","doi-asserted-by":"publisher","first-page":"7390","DOI":"10.1074\/jbc.M708192200","volume":"283","author":"F Giorgini","year":"2008","unstructured":"Giorgini F, M\u00f6ller T, Kwan W, Zwilling D, Wacker JL, Hong S, et al. Histone deacetylase inhibition modulates kynurenine pathway activation in yeast, microglia, and mice expressing a mutant huntingtin fragment. J Biol Chem. 2008;283:7390\u2013400.","journal-title":"J Biol Chem"},{"key":"1782_CR70","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/ncomms11758","volume":"7","author":"A Benraiss","year":"2016","unstructured":"Benraiss A, Wang S, Herrlinger S, Li X, Chandler-Militello D, Mauceri J, et al. Human glia can both induce and rescue aspects of disease phenotype in Huntington disease. Nat Commun. 2016;7:1\u201313.","journal-title":"Nat Commun."}],"container-title":["Journal of Neuroinflammation"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1186\/s12974-020-01782-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/article\/10.1186\/s12974-020-01782-9\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1186\/s12974-020-01782-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,4,1]],"date-time":"2021-04-01T19:08:54Z","timestamp":1617304134000},"score":1,"resource":{"primary":{"URL":"https:\/\/jneuroinflammation.biomedcentral.com\/articles\/10.1186\/s12974-020-01782-9"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,4,2]]},"references-count":70,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2020,12]]}},"alternative-id":["1782"],"URL":"https:\/\/doi.org\/10.1186\/s12974-020-01782-9","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.2.19435\/v1","asserted-by":"object"}]},"ISSN":["1742-2094"],"issn-type":[{"value":"1742-2094","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,4,2]]},"assertion":[{"value":"18 December 2019","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"20 March 2020","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 April 2020","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"All experiments were approved and performed under the guidelines of the Institutional Animal Ethics Committees, in conformity with the Canadian Council on Animal Care guidelines as administered by the Animal Care Committee of Universit\u00e9 Laval.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval"}},{"value":"Not applicable.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"98"}}