Complete List of Publications Here 


K. Lin, G. Bieri, Geraldine Gontier, S. Muller, L.K. Smith, C.E. Snethlage, C.W. White III, S. Maybury-Lewis, and S.A. Villeda. MHC Class I H2-Kb negatively regulates neural progenitor cell proliferation by inhibiting FGFR signalingPLOS Biology19(6): e3001311.  

T.T. Ho, P.V. Dellorusso, E.V. Verovskaya, S.T. Bakker, J. Flach, L.K. Smith, P.B. Ventura, O.M. Lansinger, A. Herault, S.Y. Zhang, Y-A. Kang, S.A. Villeda, and E. Passegue. Aged hematopoietic stem cells are refractory to systemic interventionsJournal of Experimental Medicine218(7):e20210223



A.M. Horowitz, X. Fan, G. Bieri, L.K. Smith, C.I. Sanchez-Diaz, A.B. Schroer, G. Gontier, K.B. Casaletto, J.H. Kramer, K.E. Williams, and S.A. Villeda. Blood factors transfer beneficial effects of exercise on neurogenesis and cognition to the aged brainScience369(6500):167-173.

  • Insights: Exercising your mind. Science. 369(6500):144-145.
  • Research Highlight: Blood from fit mice bestows brain benefits of exercise. Nature. 583:335.
  • Comment: A new player in the beneficial effects of exercise on the aged brain. Signal Transduct Target Ther. 5:184.

C.W. White III, X. Fan, J.C. Maynard, E.G. Wheatley, G. Bieri, J. Couthois, A.L. Burlingame, and S.A. Villeda. Age-related loss of neural stem cell O-GlcNAc promotes a glial cell fate switch through STAT3 activationProceeding of the National Academy of Sciences USA177(36):22214-22214.  

L.K. Smith, E. Verovskaya, G. Bieri, A.M. Horowitz, S.N.I. von Ungern-Sternberg, K. Lin, P. Seizer, E. Passegue, and S.A. Villeda. The aged hematopoietic system promotes hippocampal-dependent cognitive declineAging Cell19(8):e13192.  

D. Brigger, C. Riether, Robin van Brummelen, K.I. Mosher, A. Shiu, Z. Ding, H. Yousef, J.M. Castellano, D. Storni, N. Graff-Radford, M. Britschgi, D. Grandgirard, M. Siegrist, N. Moullan, W. Hofstetter, S.L. Leib, P.M. Villiger, J. Auwerx, S.A. Villeda, T. Wyss-Coray, M. Noti, and A. Eggel. Eosinophils regulate adipose tissue inflammation and sustain physical and immunological fitness in old ageNature Metabolism2(8):688-702.

Q-H. Nguyen, R.G. Witt, B. Wang, C. Eikani, J. Shea, L. Smith, G. Boyle, J. Cadaoas, R. Sper, J.D. MacKenzie, S. Villeda and T.C. MacKenzie. Tolerance induction and microglial engraftment after fetal therapy without conditioning in mice with Mucopolysaccharidosis Type VIIScience Translational Medicine. 12(532):8980.

J.M Shea and S.A. Villeda. Dampening the power of the brain – when aging meets cognitionJournal of Gerontology: Biological Sciences DOI:10.1093/Gerona/glaa170.



E.G. Wheatley, E. Albarran, C.W. White III, G. Bieri, C.I. Sanchez-Diaz, K. Pratt, C.E. Snethlage, J.B. Ding, and S.A. Villeda. Neuronal O-GlcNAcylation improves cognitive function in the aged mouse brainCurrent Biology29(20):3359-3369.

C. Schneider, J. Lee, S. Koga, R.R. Ricardo-Gonzalez, J.C. Nussbaum, L.K. Smith, S.A. Villeda, H.E. Liang and R.M. Locksley.  Tissue-resident group 2 innate lymphoid cells differentiate by layered ontogeny and in situ perinatal priming. Immunity. 7613(19):30199-2. 

P. Sousa-Victor, J. Neves, W. Cedron-Craft, P.B. Ventura, C-Y. Liao, R.R. Riley, I. Soifer, N. van Bruggen, G.A. Kolumam, S.A. Villeda, D.A. Lamba and H. Jasper. MANF regulates metabolic and immune homeostasis in ageing and protects against liver damage. Nature Metabolism. 1(2):276-290.

S.N. Austad, M.A. Wood, S.A Villeda, J.L. Voss, A. Sahay and M. Albert. Innovative approaches in cognitive aging. Neurobiology of Aging4580(19):30127-7.  

A.B. Schroer, A.M. Horowitz and S.A Villeda. Platelets give a running start to adult hippocampal neurogenesis. Stem Cell Reports12(4):643-646.  

C.W. White 3rd, K. Pratt, S.A. Villeda. OPCs on a Diet: A Youthful Serving of Remyelination. Cell Metabolism. 30(6):1004-1006. 



G. Gontier, M. Iyer, J.M. Shea, G. Bieri, E.G. Wheatley, M. Ramalho-Santos, and S.A. Villeda. Tet2 rescues age-related regenerative decline and enhances cognitive function in the adult mouse brainCell Reports22(8):1974-1981.

G.B. Bieri. K.M. Lucin, C.E. O’Brien, S.A. Villeda, and T. Wyss-Coray.  Proteolytic cleavage of Beclin 1 exacerbates neurodegeneration. Molecular Neurodegeneration13(1):68.

L.K. Smith, C.W. White and S.A Villeda. The systemic environment: at the interface of aging and adult neurogenesis. Cell Tissue Res. 371(1):105-113.  



A. Deczkowska, O. Matcovitch-Natan, A. Tsitsou-Kampeli, S. Ben-Hamo, R. Dvir-Szternfeld, A. Spinrad, O. Singer, E. David, D.R. Winter, L.K. Smith, A. Kertser, K. Baruch, N. Rosenzweig, A. Terem, M. Prinz, S. Villeda, A. Citri, I. Amit, and M. Schwartz. Mef2C restrains microglial inflammatory response and is lost in brain ageing in an INF-I-dependent manner. Nature Communications. 8(1):717. 

X. Fan, E.G. Wheatley and S.A Villeda. Mechanisms of hippocampal aging and the potential for rejuvenation. Annual Review of Neuroscience40: 251-271.  

A.M. Horowitz and S.A Villeda. Therapeutic potential of systemic brain rejuvenation strategies for neurodegenerative disease. F1000 ResearchDOI: 10.1007/s00441-017-2715-8.



Y. Zhang, T.L. Roth, E.E. Gray, H. Chen, L.B. Rodda, Y. Liang, P. Ventura, S. Villeda, P.R. Crocker, and J.G. Cyster. Mechanisms of innate-like lymphocyte surveillance of the subcapsular sinus to provide lymph node barrier immunity. eLifeDOI: 10.7554/3Life.18156  

J. Middeldorp, B. Lehallier, S.A. Villeda, S. Miedema, E. Evans, E. Czirr, H. Zhang, J. Luo, T. Stan, K. Mosher, E. Masliah and T. Wyss-Coray. Pre-clinical assessment of young plasma for Alzheimer’s disease. JAMA Neurology. 73(11):1325-1333.

J. U. Udeochu, J. M. Shea, and S.A. Villeda. Microglia communication: Parallels between aging and Alzheimer’s disease. Clinical and Experimental Neuroimmunology7(2):114-125.  



L.K. Smith, Y.B. He, J.S. Park, G. Bieri, C. Snethlage, K. Lin, G. Gontier, R. Wabl, K. Plambeck, J. Udeochu, E.G. Wheatley, J. Bouchard, A. Eggel, R. Narasimha, J.L. Grant, J. Lou, T. Wyss-Coray and S.A. Villedaβ2-Microglobulin is a systemic pro-aging factor that impairs cognitive function and neurogenesisNature Medicine21(8):932-37. 

  • News and Views: Breaking bad blood. Nature Medicine. 21(8):844-45
  • Highlight: ‘Old-age protein’ may cause memory loss. ScienceNow. AAAS; 2015

M. Armanios, R. De Cabo, J. Mannick, L. Partridge, J. Van Deursen, and S. Villeda. Translational strategies in aging and age-related disease. Nature Medicine. 21(12):1-5.  

G. Gontier, K. Lin and S.A. Villeda. Fat chance for neural stem cells in Alzheimer’s disease. Cell Stem Cell. 17(4):373-374.  



S.A. Villeda*, K. Plambeck, J. Middeldorp, J.M. Castellano, K.I. Mosher, J. Luo, L.K. Smith, G. Bieri, K. Lin, D. Berdnik, R. Wabl, J. Udeochu, E.G. Wheatley, B. Zou, D. Simmons, X.S. Xie, F. Longo, T. Wyss-Coray*. Young blood reverses age-related impairments in cognitive function and synaptic plasticity in miceNature Medicine. 20(6):659-63 *co-corresponding author

  • News and Views: Young blood rejuvenates old brains. Nature Medicine. 20(6):582-83
  • Highlight: Young Blood. New England Journal of Medicine371(6):573-5
  • Highlight: Blood-derived rejuvenation. Nat. Rev. Neurosci. 15:352
  • Highlight: Could young blood combat age-related cognitive decline. Nat. Rev. Neurol. 
  • Rejuvenation: It’s in our blood. Cell Metabolism20(1):2-4
  • Highlight: Young blood renews old mice. ScienceNow. AAAS; 2014

J. Bouchard & S.A. Villeda. Aging and Brain Rejuvenation as Systemic Events. Journal of Neurochemistry. 132(1):5-19.



S.A. Villeda, J. Luo, K.I. Mosher, B. Zou, M. Britschgi, G. Bieri, T.M. Stan, N. Fainberg,  Z. Ding, A. Eggel, K.M. Lucin, E. Czirr, J-S. Park, S. Doupres-Couillard, L. Aigner, G. Li, E.R. Peskind, J.A. Kaye, J.F. Quinn, D.R. Galasko, X.S. Xie, T.A. Rando, and T. Wyss-Coray. The aging systemic milieu negatively regulates neurogenesis and cognitive functionNature. 477(7362):90-4  

  • News and Views: Ageing: Blood Ties. Nature. 477(7362):41-2
  • Highlight: Ageing, it’s in the blood. Nat. Rev. Neurosci. 12(10):547
  • Highlight: The Brain’s Fountain of Youth. ScienceNow. AAAS; 2011