[1]

Shea JM, Villeda SA. 2025. Microglia aging in the hippocampus advances through intermediate states that drive activation and cognitive decline. eLife 13:RP97671

doi: 10.7554/eLife.97671.3
[2]

Galatro TF, Holtman IR, Lerario AM, Vainchtein ID, Brouwer N, et al. 2017. Transcriptomic analysis of purified human cortical microglia reveals age-associated changes. Nature Neuroscience 20:1162−1171

doi: 10.1038/nn.4597
[3]

Li X, Li Y, Jin Y, Zhang Y, Wu J, et al. 2023. Transcriptional and epigenetic decoding of the microglial aging process. Nature Aging 3:1288−1311

doi: 10.1038/s43587-023-00479-x
[4]

Carr L, Mustafa S, Collins-Praino LE. 2025. The hallmarks of ageing in microglia. Cellular and Molecular Neurobiology 45:45

doi: 10.1007/s10571-025-01564-y
[5]

Ma F, Sen R. 2026. Physiological aging in three dimensions. Trends in Cell Biology 36:230−245

doi: 10.1016/j.tcb.2025.08.002
[6]

Keren-Shaul H, Spinrad A, Weiner A, Matcovitch-Natan O, Dvir-Szternfeld R, et al. 2017. A unique microglia type associated with restricting development of Alzheimer's disease. Cell 169:1276−1290.e17

doi: 10.1016/j.cell.2017.05.018
[7]

Huang Z, Merrihew GE, Larson EB, Park J, Plubell D, et al. 2023. Brain proteomic analysis implicates actin filament processes and injury response in resilience to Alzheimer's disease. Nature Communications 14:2747

doi: 10.1038/s41467-023-38376-x
[8]

Baligács N, Albertini G, Borrie SC, Serneels L, Pridans C, et al. 2024. Homeostatic microglia initially seed and activated microglia later reshape amyloid plaques in Alzheimer's Disease. Nature Communications 15:10634

doi: 10.1038/s41467-024-54779-w
[9]

Uhlemann R, Gertz K, Boehmerle W, Schwarz T, Nolte C, et al. 2016. Actin dynamics shape microglia effector functions. Brain Structure and Function 221:2717−2734

doi: 10.1007/s00429-015-1067-y
[10]

Bajpai A, Li R, Chen W. 2021. The cellular mechanobiology of aging: from biology to mechanics. Annals of the New York Academy of Sciences 1491:3−24

doi: 10.1111/nyas.14529
[11]

Portugal CC, Almeida TO, Tedim-Moreira J, Silva C, Canedo T, et al. 2025. Profilin 1 controls a microglial cytoskeleton checkpoint to prevent senescence and premature synaptic decline. Journal of Neuroinflammation 22:264

doi: 10.1186/s12974-025-03588-z
[12]

Sanchini C, Rosito M, Bartolini F, Di Angelantonio S. 2026. Targeting microglia microtubules: cytoskeletal remodeling as a druggable hub in neuroinflammation and neurodegeneration. Frontiers in Neuroscience 20:1812417

doi: 10.3389/fnins.2026.1812417
[13]

Kuhn J, Banerjee P, Haye A, Robinson DN, Iglesias PA, et al. 2025. Complementary cytoskeletal feedback loops control signal transduction excitability and cell polarity. Nature Communications 16:7482

doi: 10.1038/s41467-025-62799-3
[14]

Adrian M, Weber M, Tsai MC, Glock C, Kahn OI, et al. 2023. Polarized microtubule remodeling transforms the morphology of reactive microglia and drives cytokine release. Nature Communications 14:6322

doi: 10.1038/s41467-023-41891-6
[15]

Rosito M, Sanchini C, Gosti G, Moreno M, De Panfilis S, et al. 2023. Microglia reactivity entails microtubule remodeling from acentrosomal to centrosomal arrays. Cell Reports 42:112104

doi: 10.1016/j.celrep.2023.112104
[16]

Hu J, Chen Q, Zhu H, Hou L, Liu W, et al. 2023. Microglial Piezo1 senses Aβ fibril stiffness to restrict Alzheimer's disease. Neuron 111:15−29.e8

doi: 10.1016/j.neuron.2022.10.021
[17]

Badimon A, Strasburger HJ, Ayata P, Chen X, Nair A, et al. 2020. Negative feedback control of neuronal activity by microglia. Nature 586:417−423

doi: 10.1038/s41586-020-2777-8
[18]

Khodaee F, Zandie R, Leger LA, Xia Y, Thadawasin P, et al. 2025. The dissipation theory of aging: a quantitative analysis using a cellular aging map. npj Aging 11:86

doi: 10.1038/s41514-025-00277-2
[19]

Bussian TJ, Aziz A, Meyer CF, Swenson BL, van Deursen JM, et al. 2018. Clearance of senescent glial cells prevents tau-dependent pathology and cognitive decline. Nature 562:578−582

doi: 10.1038/s41586-018-0543-y
[20]

Elmore MRP, Hohsfield LA, Kramár EA, Soreq L, Lee RJ, et al. 2018. Replacement of microglia in the aged brain reverses cognitive, synaptic, and neuronal deficits in mice. Aging Cell 17:e12832

doi: 10.1111/acel.12832
[21]

Guneykaya D, Ivanov A, Hernandez DP, Haage V, Wojtas B, et al. 2018. Transcriptional and translational differences of microglia from male and female brains. Cell Reports 24:2773−2783.e6

doi: 10.1016/j.celrep.2018.08.001
[22]

Villa A, Gelosa P, Castiglioni L, Cimino M, Rizzi N, et al. 2018. Sex-specific features of microglia from adult mice. Cell Reports 23:3501−3511

doi: 10.1016/j.celrep.2018.05.048
[23]

Kang S, Ko EY, Andrews AE, Shin JE, Nance KJ, et al. 2024. Microglia undergo sex-dimorphic transcriptional and metabolic rewiring during aging. Journal of Neuroinflammation 21:150

doi: 10.1186/s12974-024-03130-7
[24]

Seto M, Clifton M, Gomez ML, Coughlan G, Gifford KA, et al. 2025. Sex-specific associations of gene expression with Alzheimer's disease neuropathology and ante-mortem cognitive performance. Nature Communications 16:9466

doi: 10.1038/s41467-025-64525-5
[25]

Nebel RA, Aggarwal NT, Barnes LL, Gallagher A, Goldstein JM, et al. 2018. Understanding the impact of sex and gender in Alzheimer's disease: a call to action. Alzheimer's & Dementia 14:1171−1183

doi: 10.1016/j.jalz.2018.04.008
[26]

Socodato R, Portugal CC, Canedo T, Rodrigues A, Almeida TO, et al. 2020. Microglia dysfunction caused by the loss of rhoa disrupts neuronal physiology and leads to neurodegeneration. Cell Reports 31:107796

doi: 10.1016/j.celrep.2020.107796
[27]

Socodato R, Rodrigues-Santos A, Tedim-Moreira J, Almeida TO, Canedo T, et al. 2023. RhoA balances microglial reactivity and survival during neuroinflammation. Cell Death & Disease 14:690

doi: 10.1038/s41419-023-06217-w
[28]

Socodato R, Almeida TO, Portugal CC, Santos ECS, Tedim-Moreira J, et al. 2023. Microglial Rac1 is essential for experience-dependent brain plasticity and cognitive performance. Cell Reports 42:113447

doi: 10.1016/j.celrep.2023.113447
[29]

Bokoch GM. 2003. Biology of the p21-activated kinases. Annual Review of Biochemistry 72:743−781

doi: 10.1146/annurev.biochem.72.121801.161742
[30]

Bernstein BW, Bamburg JR. 2010. ADF/Cofilin: a functional node in cell biology. Trends in Cell Biology 20:187−195

doi: 10.1016/j.tcb.2010.01.001
[31]

Arani A, Murphy MC, Glaser KJ, Manduca A, Lake DS, et al. 2015. Measuring the effects of aging and sex on regional brain stiffness with MR elastography in healthy older adults. NeuroImage 111:59−64

doi: 10.1016/j.neuroimage.2015.02.016
[32]

Cook M, Lin H, Mishra SK, Wang GY. 2022. BAY 11-7082 inhibits the secretion of interleukin-6 by senescent human microglia. Biochemical and Biophysical Research Communications 617:30−35

doi: 10.1016/j.bbrc.2022.05.090
[33]

Kessels S, Trippaers C, Mertens M, Hamad I, Rombaut B, et al. 2025. Cytoskeletal control in adult microglia is essential to restore neurodevelopmental synaptic and cognitive deficits. Science Advances 11:eadw0128

doi: 10.1126/sciadv.adw0128
[34]

Crapser JD, Spangenberg EE, Barahona RA, Arreola MA, Hohsfield LA, et al. 2020. Microglia facilitate loss of perineuronal nets in the Alzheimer's disease brain. EBioMedicine 58:102919

doi: 10.1016/j.ebiom.2020.102919
[35]

Cabungcal JH, Steullet P, Morishita H, Kraftsik R, Cuenod M, et al. 2013. Perineuronal nets protect fast-spiking interneurons against oxidative stress. Proceedings of the National Academy of Sciences of the United States of America 110:9130−9135

doi: 10.1073/pnas.1300454110
[36]

Murthy SE, Dubin AE, Patapoutian A. 2017. Piezos thrive under pressure: mechanically activated ion channels in health and disease. Nature Reviews Molecular Cell Biology 18:771−783

doi: 10.1038/nrm.2017.92
[37]

Sell DR, Monnier VM. 2012. Molecular basis of arterial stiffening: role of glycation – a mini-review. Gerontology 58:227−237

doi: 10.1159/000334668
[38]

Baker AM, Bird D, Lang G, Cox TR, Erler JT. 2013. Lysyl oxidase enzymatic function increases stiffness to drive colorectal cancer progression through FAK. Oncogene 32:1863−1868

doi: 10.1038/onc.2012.202
[39]

Tarumi T, Khan MA, Liu J, Tseng BM, Parker R, et al. 2014. Cerebral hemodynamics in normal aging: central artery stiffness, wave reflection, and pressure pulsatility. Journal of Cerebral Blood Flow & Metabolism 34:971−978

doi: 10.1038/jcbfm.2014.44
[40]

Silver J, Miller JH. 2004. Regeneration beyond the glial scar. Nature Reviews Neuroscience 5:146−156

doi: 10.1038/nrn1326
[41]

Zheng Q, Liu H, Yu W, Dong Y, Zhou L, et al. 2023. Mechanical properties of the brain: focus on the essential role of Piezo1-mediated mechanotransduction in the CNS. Brain and Behavior 13:e3136

doi: 10.1002/brb3.3136
[42]

Viji Babu PK, Radmacher M. 2019. Mechanics of brain tissues studied by atomic force microscopy: a perspective. Frontiers in Neuroscience 13:600

doi: 10.3389/fnins.2019.00600
[43]

Jorstad NL, Song JHT, Exposito-Alonso D, Suresh H, Castro-Pacheco N, et al. 2023. Comparative transcriptomics reveals human-specific cortical features. Science 382:eade9516

doi: 10.1126/science.ade9516
[44]

Gabitto MI, Travaglini KJ, Rachleff VM, Kaplan ES, Long B, et al. 2024. Integrated multimodal cell atlas of Alzheimer's disease. Nature Neuroscience 27:2366−2383

doi: 10.1038/s41593-024-01774-5
[45]

Olah M, Patrick E, Villani AC, Xu J, White CC, et al. 2018. A transcriptomic atlas of aged human microglia. Nature Communications 9:539

doi: 10.1038/s41467-018-02926-5
[46]

Hammond TR, Dufort C, Dissing-Olesen L, Giera S, Young A, et al. 2019. Single-cell RNA sequencing of microglia throughout the mouse lifespan and in the injured brain reveals complex cell-state changes. Immunity 50:253−271.e6

doi: 10.1016/j.immuni.2018.11.004
[47]

Pollard TD. 1986. Rate constants for the reactions of ATP- and ADP-actin with the ends of actin filaments. The Journal of Cell Biology 103:2747−2754

doi: 10.1083/jcb.103.6.2747
[48]

Kuhn JR, Pollard TD. 2005. Real-time measurements of actin filament polymerization by total internal reflection fluorescence microscopy. Biophysical Journal 88:1387−1402

doi: 10.1529/biophysj.104.047399
[49]

Schafer DA, Jennings PB, Cooper JA. 1996. Dynamics of capping protein and actin assembly in vitro: uncapping barbed ends by polyphosphoinositides. The Journal of Cell Biology 135:169−179

doi: 10.1083/jcb.135.1.169
[50]

Mullins RD, Heuser JA, Pollard TD. 1998. The interaction of Arp2/3 complex with actin: nucleation, high affinity pointed end capping, and formation of branching networks of filaments. Proceedings of the National Academy of Sciences of the United States of America 95:6181−6186

doi: 10.1073/pnas.95.11.6181
[51]

Pantaloni D, Carlier MF. 1993. How profilin promotes actin filament assembly in the presence of thymosin β4. Cell 75:1007−1014

doi: 10.1016/0092-8674(93)90544-Z