Guo, NN Cover
Guo, 2011
Smrt cover
Smrt, 2007

(Authors: $:graduate students; %:undergrad students/research interns)


  • Gao Y, Syed M%, and Zhao X. Mechanisms underlying the effect of voluntary running on adult hippocampal neurogenesis. Hippocampus 2023  Mar 9. PMID: 36892196 DOI: 10.1002/hipo.23520
  • S. RosasK. A. Schoeller%, E. Chang%, H. MeiM. A. KatsK. W. EliceiriX. ZhaoF. Yesilkoy Metasurface-enhanced mid-infrared spectrochemical imaging of tissues (under review) (preprint: 2023 (In this collaborative paper, Keegan and Edward in our lab generated and optimized mouse brain tissue processing)
  • Chenfeng He, Noah Cohen Kalafut, Soraya O. Sandoval$, Ryan Risgaard$, Carissa L. Sirois, Chen Yang, Saniya Khullar$, Marin Suzuki, Xiang Huang, Qiang Chang, Xinyu Zhao Andre M.M. Sousa, Daifeng Wang:  Brain and Organoid Manifold Alignment (BOMA) reveals cell-type developmental functional genomics across in-vivo brains and in-vitro organoids. Cell Reports Methods 2023 ( (news release) (In this collaborative paper, Soraya and Carissa in our lab generated and analyzed human forebrain organoids to compare with human brains as a validation of BOMA computation method))
  • Ludwig AL$, Mayerl SJ$, Gao Y, Banghart M, Bacig C, Fernandez Zepeda MA, Zhao X, Gamm DM. Re-formation of synaptic connectivity in dissociated human stem cell-derived retinal organoid cultures.Proc Natl Acad Sci U S A. 2023 Jan 10;120(2):e2213418120. doi: 10.1073/pnas.2213418120. Epub 2023 (In this collaborative paper, Yu Gao in our lab created all the rabies virus for monosynaptic tracing and lentivirus targeting starter cells to compare with human brains as a validation of BOMA computation method)


  • Canales A$, Scheuer KS$, Zhao X, Jackson MB. Unitary synaptic responses of parvalbumin interneurons evoked by excitatory neurons in the mouse barrel cortex. Cereb Cortex. 2022 Oct 13;. doi: 10.1093/cercor/bhac403. [Epub ahead of print] PubMed PMID: 36227216.
  • Sahar Javadi$, Yue Li, Jie Sheng%, Lucy Zhao%, Yao Fu%,  Daifeng Wang, and Xinyu Zhao. Sustained correction of hippocampal neurogenic and cognitive deficits after a brief treatment by Nutlin-3 in a mouse model of Fragile X Syndrome. BMC Med 2022 May 13;20(1):163.doi: 10.1186/s12916-022-02370-9. (news release)
  • Hunt JFV$ , Li M, Risgaard RD%,  Ananiev GE, Wildman S, Zhang F, Bugni TS, Zhao X,* and Bhattacharyya  A* (*co-correspondance)  High Throughput Small Molecule Screen for Reactivation of FMR1 in Fragile X Syndrome Human Neural Cells, Cells. 2022 Jan; 11(1): 69. Published online 2021 Dec 27. doi: 10.3390/cells11010069
  • Shen M, Guo Y, Dong Q, Gao, Y, Stockton ME%, Li M, Kannan S%, Korabelnikov T%, Schoeller, AK%, Sioris CL, Zhou C$, Le J%, Chang C,  Sun, Q-Q, Wang D, and Zhao X. FXR1 regulation of parvalbumin interneurons in the prefrontal cortex is critical for schizophrenia-like behaviors. Mol Psychiatry 2021 Apr 16. doi: 10.1038/s41380-021-01096-z. PMID: 33863995
  • Joel Richter and Xinyu Zhao: The molecular biology of FMRP: new insights into fragile X syndrome. Nature Review Neuroscience 2021 Feb 19.  PMID: 33608673; doi: 10.1038/s41583-021-00432-0. ( link)



  • Jichao Sun, Jared Carlson-Stevermer, Utpal Das, Minjie Shen, Marion Delenclos, Amanda Snead, Soo Koo, Lina Wang, Dianhua Qiao, Jonathan Loi, ANDREW PETERSEN, Michael Stockton, Anita Bhattacharyya, Mathew Jones, Xinyu Zhao, Pam McLean, Andrew Sproul, Krishanu Saha, and Subhojit Roy. CRISPR/Cas9 editing of APP C-terminus attenuates b-cleavage and promotes a-cleavage” Nature Communications (in press) [#NCOMMS-18-25145C]
  • Zhao# and Bhattacharyya# (#co-correspondence). “The need for human models to study human neurodevelopmental disorders American Journal of Human Genetics, (2018)
  • Yue Li*, Minjie Shen* (*equal contribution), Michael Stockton: “Hippocampal deficits in neurodevelopmental disorders” Neurobiology of Learning and Memory, 2018
  • Liu B*, Li Y*, (*equal contribution), Stackpole, Novak A, Gao Y, Zhao Y, Zhao X#, Richter JD# (#co-correspondence).  Regulatory Discrimination of mRNAs by FMRP Controls Mouse Adult Neural Stem Cell DifferentiationPNAS 2018
    Faculty 1000 recommended exceptional.
  • Mo, K., Wu, S., Gu, X., Xiong, M., Cai, W., Atianjoh, F.E., Jobe, E.E$., Zhao, X., Tu, W.F., and Tao, Y.X. (2018). MBD1 contributes to the genesis of acute pain and neuropathic pain by epigenetic silencing of Oprm1 and Kcna2 genes in primary sensory neurons. J Neurosci. 2018 Nov 14;38(46):9883-9899. doi: 10.1523/JNEUROSCI.0880-18.2018. Epub 2018 Sep 28. PubMed PMID: 30266739; PubMed Central PMCID: PMC6234297
  • Sun, Y*., Gao, Y*., (*equal contribution), Tidei%, J.J., Shen, M., Hoang, J.T.%, Wagner, D.%, and Zhao, X. (2018). Loss of MeCP2 in immature neurons leads to impaired network integration. Hum Mol Genet. (in press) see preprint 
  • Li Y., Stockton ME%, Eisinger BE, Zhao Y, Miller JL%, Bhuiyan I.%, Gao Y, Wu Z, Peng J, and Zhao X, Reducing histone acetylation rescues cognitive deficits in a mouse model of Fragile X syndrome. 2018 Nature Communications 9, 2494. June 27 2018. (new release)
  • Hunt JFV$, Li M, Zhao X*, Bhattacharyya A* (*co-correspondence). Using human neural progenitor cell models to conduct large-scale drug screens for neurological and psychiatric diseases (MiMB chapter 2018)
  • Li M, Hunt J$, Bhattacharyya A*. Zhao X* (*co-correspondence), One-step generation of seamless luciferase gene knock-in using CRISPR/Cas9 genome editing in human pluripotent stem cells (MiMB chapter 2018)
  • Patzlaff NP$, Shen M, Zhao X. Regulation of Adult Neurogenesis by the Fragile X Family of RNA Binding Proteins Brain Plasticity 2018 Aug 10;3(2):205-223. doi: 10.3233/BPL-170061. Review. PubMed PMID: 30151344; PubMed Central PMCID: PMC6091053.
  • Kempermann G, Gage FH, Aigner L, Song H, Curtis MA, Thuret S, Kuhn HG, Jessberger S, Frankland PW, Cameron HA, Gould E, Hen R, Abrous DN, Toni N, Schinder AF, Zhao X, Lucassen PJ, Frisén J. Human Adult Neurogenesis: Evidence and Remaining Questions. Cell Stem Cell. 2018 Apr 18. pii: S1934-5909(18)30166-8. doi: 10.1016/j.stem.2018.04.004. [Epub ahead of print]
  • Eisinger BE, Zhao X. Identifying molecular mediators of environmentally enhanced neurogenesis. Cell Tissue Res. 2018 Jan;371(1):7-21. doi: 10.1007/s00441-017-2718-5. Epub 2017 Nov 10. Review. PubMed PMID: 29127518.
  • Zhao X, Moore DL. Neural stem cells: developmental mechanisms and disease modeling.Cell Tissue Res. 2018 Jan;371(1):1-6. doi: 10.1007/s00441-017-2738-1. PubMed PMID: 29196810.


  • Zhang P, Rausch C, Hastert FD, Boneva B, Filatova A, Patil SJ, Nuber UA, Gao Y, Zhao X, Cardoso MC. Methyl-CpG binding domain protein 1 regulates localization and activity of Tet1 in a CXXC3 domain-dependent manner. (2017) Nucleic Acids Res.Apr 25. doi: 10.1093/nar/gkx281. [Epub ahead of print] PMID: 28449087
  • Patzlaff NP$, Nemec KM%, Malone S%, Li Y, Zhao X. Fragile X related protein 1 (FXR1P) regulates proliferation of adult neural stem cells. (2017) Hum Mol Genet  Apr 1;26(7):1340-1352. doi: 10.1093/hmg/ddx034.  Fragile X related protein 1 (FXR1P) regulates proliferation of adult neural stem cells. PMID: 28204491
  • Sun L, Zhao JY, Gu X, Liang L, Wu S, Mo K, Feng J, Guo W, Zhang J, Bekker A, Zhao X, Nestler EJ, Tao YX. Nerve injury-induced epigenetic silencing of opioid receptors controlled by DNMT3a in primary afferent neurons.(2017) Pain. 2017 Mar 4. doi: 10.1097/j.pain.0000000000000894. [Epub ahead of print] PMID:28267064
  • Jobe EM$*, Gao Y* (*equal contribution), Eisinger BE, Mladucky JK%, Giuliani CC%, Kelnhofer LE%, and Zhao X. Methyl-CpG binding protein MBD1 regulates neuronal lineage commitment through maintaining adult neural stem cell identity (2017). Journal of Neuroscience. Jan 18;37(3):523-536. doi: 10.1523/JNEUROSCI.  PMID:26989163. (see news release). see a featured Journal Club discussing the impact of this paper on JN website.
  • Gao Y*, Wang F*, Eisinger BE*, Kelnhofer LE%, Jobe EM$, Zhao X. “Integrative Single-Cell Transcriptomics Reveal Molecular Networks Defining Neuronal Maturation during Postnatal Neurogenesis.” (2017) Cerebral Cortex. Mar 1;27(3):2064-2077. doi: 10.1093/cercor/bhw040. PMID:26989163. [* contributed equally (see news release)]


  • Wang F, Tidei JJ%, Polich ED%, Gao Y,  Zhao H, Perrone-Bizzozero NI, Weixiang Guo , Zhao X. Positive feedback between RNA-binding protein HuD and transcription factor SATB1 promotes neurogenesis. Proc Natl Acad Sci U S A plus (2015)
  • Guo W, Polich ED%, Juan Su, J, Gao Y, Christopher, DM%, Allan AM, Wang M, Wang F, Wang G, Zhao X. Fragile X Proteins FMRP and FXR2P control synaptic GluA1 expression and neuronal maturation via distinct mechanisms. Cell Reports. 2015. NIHMS690530, Publ.ID: CELREP1788
  • Gao Y*, Su J* (*,equal contributions), Guo W, Polich ED%, Magyar DP%, Xing Y, Li H$, Smrt RD$, Chang Q, Zhao X. Inhibition of miR-15a Promotes BDNF Expression and Rescues Dendritic Maturation Deficits in MeCP2-Deficient Neurons. Stem Cells. 2015 May;33(5):1618-29. PubMed PMID: 25639236; PubMed Central PMCID: PMC4409556.
  • Guo W, Tsujimura K, Otsuka I M, Irie K, Igarashi K, Nakashima K, Zhao X. VPA alleviates neurological deficits and restores gene expression in a mouse model of Rett syndrome. PLoS One. 2014;9(6):e100215. PubMed PMID: 24968028; PubMed Central PMCID: PMC4072629.
  • Li H$, Zhong X, Chau KF, Santistevan NJ%, Guo W, Kong G, Li X, Kadakia M, Masliah J, Chi J, Jin P, Zhang J, Zhao X, Chang Q. Cell cycle-linked MeCP2 phosphorylation modulates adult neurogenesis involving the Notch signalling pathway. Nature Communications. 2014 Nov 25;5:5601. PubMed PMID: 25420914; PubMed Central PMCID: PMC4288926.
  • Chow CL$, Guo W, Trivedi P, Zhao X, Gubbels SP.Characterization of a unique cell population marked by transgene expression in the adult cochlea of nestin-CreER(T2) /tdTomato-reporter mice. Journal of Comparative Neurology. 2015 Jun 1;523(10):1474-87. doi: 10.1002/cne.23747
  • Li Y, Zhao X.Concise review: Fragile X proteins in stem cell maintenance and differentiation. Stem Cells. 2014 Jul;32(7):1724-33. doi: 10.1002/stem.1698. (Concise Review). PubMed PMID: 24648324; PubMed Central PMCID: PMC4255947.
  • Hsieh J and Zhao X “Adult Neurogenesis: Genetics and Epigenetics in Adult Neuorgenesis” Cold Spring Harbor Laboratory Press (2015, book chapter)
  • Guo W and Zhao X. “Differential intrinsic and extrinsic regulations of the two Neurogenic Niches in the adult brain”.  Stem Cell Biology and Regenerative Medicine (Springer, Series Editor, Kursad Turksen) (2015 )






Before 2003 (Zhao Lab started in 2003)

(see a complete list of published work by Dr. Zhao:  )