Howard Gritton's directory photo.

Howard Gritton

Assistant Professor

Primary Affiliation

Neurotechnology for Memory and Cognition


Status Affiliate Faculty

Home Department of Comparative Biosciences




  • Biography

    Howard Gritton is an assistant professor in the Department of Comparative Biosciences. His primary affiliation is Neurotechnology for Memory and Cognition.


    • B.S., biology, University of Utah, Salt Lake City, Utah

    • Ph.D., neuroscience, University of Michigan, Ann Arbor, Michigan

    • Post-doctoral training, Boston University, Boston, Massachusetts 

  • Research

    Research Areas:

    • Systems biology

    • Cognition

    • Learning and Memory

    • Acetylcholine

    • Sleep and Circadian Rhythms

    Research Interests:

    • Neuromodulation of learning and memory

    • Experience-dependent synaptic plasticity

    • Circuit mechanisms of neuropsychiatric disorders

    • Oscillatory activity in cognitive function

    • Excitatory/inhibitory balance in circuit function

    Gritton’s lab uses animal models to understand how neuromodulators contribute to sensory and memory encoding and how this can facilitate interactions across regions of the brain. They utilize optogenetic and chemogenetic targeting tools to measure how perturbations of neurotransmitter systems associated with illness or disease alter sensory encoding and the coupling of neurons and brain regions together to ultimately influence learning and memory or cognitive and social behavior in mice.

  • 2023

    • Lowet, Eric; Sheehan, Daniel J; Chialva, Ulises; De Oliveira Pena, Rodrigo; Mount, Rebecca A; Xiao, Sheng; Zhou, Samuel L; Tseng, Hua-an; Gritton, Howard; Shroff, Sanaya; Kondabolu, Krishnakanth; Cheung, Cyrus; Wang, Yangyang; Piatkevich, Kiryl D; Boyden, Edward S; Mertz, Jerome; Hasselmo, Michael E; Rotstein, Horacio G; Han, Xue ISSN: 2211-1247; DOI: 10.1016/j.celrep.2023.112906 Cell reports. , 2023, Vol.42(8), p.112906
    • Nocon, J.C., Gritton, H.J., James, N.M. et al. Parvalbumin neurons enhance temporal coding and reduce cortical noise in complex auditory scenes. Commun Biol 6, 751 (2023).
    • Shroff, S.N., Lowet, E., Sridhar, S. et al. Striatal cholinergic interneuron membrane voltage tracks locomotor rhythms in mice. Nat Commun 14, 3802 (2023).


    • Auerbach, B. D., & Gritton, H. J. (2022). Hearing in Complex Environments: Auditory Gain Control, Attention, and Hearing Loss. Frontiers in Neuroscience, 16, [799787].
    • Xiao, Sheng; Lowet, Eric; Gritton, Howard; Fabris, Pierre; Mertz, Jerome; Han, Xue; Luo, Qingming; Ding, Jun; Fu, Ling ISBN: 9781510652309; DOI: 10.1117/12.2609095 Neural Imaging and Sensing 2022 , 2022, p.5
    • Zemel, D., Gritton, H., Cheung, C., Shankar, S., Kramer, M., & Han, X. (2022). Dopamine depletion selectively disrupts interactions between striatal neuron subtypes and LFP oscillations. Cell Reports, 38(3), [110265].
    • Zemel, Dana; Gritton, Howard; Cheung, Cyrus; Shankar, Sneha; Kramer, Mark; Han, Xue ISSN: 2211-1247; DOI: 10.1016/j.celrep.2021.110265 Cell reports. , 2022, Vol.38(3), p.110265


    • Jean-Marc Tsang, Howard J. Gritton, Shoshana L. Das, Timothy D. Weber, Christopher S. Chen, Xue Han, and Jerome Mertz, "Fast, multiplane line-scan confocal microscopy using axially distributed slits," Biomed. Opt. Express 12, 1339-1350 (2021)
    • Mount, R. A., Sridhar, S., Hansen, K. R., Mohammed, A. I., Abdulkerim, M., Kessel, R., Nazer, B., Gritton, H. J., & Han, X. (2021). Distinct neuronal populations contribute to trace conditioning and extinction learning in the hippocampal CA1. eLife, 10, [e56491].
    • Mount, Rebecca A; Sridhar, Sudiksha; Hansen, Kyle R; Mohammed, Ali I; Abdulkerim, Moona E; Kessel, Robb; Nazer, Bobak; Gritton, Howard J; Han, Xue ISSN: 2050-084X; DOI: 10.7554/eLife.74730 ELife. , 2021, Vol.10, p.e74730
    • Tsang, J. M., Gritton, H. J., Das, S. L., Weber, T. D., Chen, C. S., Han, X., & Mertz, J. (2021). Fast, multiplane line-scan confocal microscopy using axially distributed slits. Biomedical Optics Express, 12(3), 1339-1350.
    • Tseng, H. A., Sherman, J., Bortz, E., Mohammed, A., Gritton, H. J., Bensussen, S., Tang, R. P., Zemel, D., Szabo, T., & Han, X. (2021). Region-specific effects of ultrasound on individual neurons in the awake mammalian brain. iScience, 24(9), [102955].
    • Tseng, Hua-An; Sherman, Jack; Bortz, Emma; Mohammed, Ali; Gritton, Howard J; Bensussen, Seth; Tang, Rockwell P; Zemel, Dana; Szabo, Thomas; Han, Xue ISSN: 2589-0042 , 2589-0042; DOI: 10.1016/j.isci.2021.102955; PMID: 34458703 iScience. , 2021, Vol.24(9), p.102955
    • Xiao, Sheng; Gritton, Howard; Tseng, Hua-an; Zemel, Dana; Han, Xue; Mertz, Jerome; Brown, Thomas G; Wilson, Tony; Waller, Laura ISSN: 0143-0343 , 1461-7374; ISBN: 9781510641334 , 9781510641341; DOI: 10.1117/12.2581250 School psychology international. , 2021, Vol.11649, p.23
    • Xiao, Sheng; Lowet, Eric; Gritton, Howard J; Fabris, Pierre; Wang, Yangyang; Sherman, Jack; Mount, Rebecca A; Tseng, Hua-An; Man, Heng-Ye; Straub, Christoph; Piatkevich, Kiryl D; Boyden, Edward S; Mertz, Jerome; Han, Xue ISSN: 2589-0042 , 2589-0042; DOI: 10.1016/j.isci.2021.103263; PMID: 34761183 iScience. , 2021, Vol.24(11), p.103263
    • Yang Y, Gritton H, Sarter M, Aton SJ, Booth V, Zochowski M (2021) Theta-gamma coupling emerges from spatially heterogeneous cholinergic neuromodulation. PLoS Comput Biol 17(7): e1009235.


    • Bensussen, Seth; Shankar, Sneha; Ching, Kimberley H; Zemel, Dana; Ta, Tina L; Mount, Rebecca A; Shroff, Sanaya N; Gritton, Howard J; Fabris, Pierre; Vanbenschoten, Hannah; Beck, Connor; Man, Heng-Ye; Han, Xue ISSN: 2589-0042 , 2589-0042; DOI: 10.1016/j.isci.2020.101330; PMID: 32674057 iScience. , 2020, Vol.23(7), p.101330
    • Sheng Xiao, Howard Gritton, Hua-An Tseng, Dana Zemel, Xue Han, and Jerome Mertz, "High-contrast multifocus microscopy with a single camera and z-splitter prism," Optica 7, 1477-1486 (2020)


    • Badon, Amaury et al. “Video-Rate Large-Scale Imaging with Multi-Z Confocal Microscopy.” Optica 6.4 (2019): n. pag. Web.
    • Badon, Amaury; Bensussen, Seth; Awal, Mehraj R; Gritton, Howard J; Gabel, Christopher V; Han, Xue; Mertz, Jerome; Goda, Keisuke; Tsia, Kevin K ISSN: 0143-0343 , 1461-7374; ISBN: 9781510624207 , 9781510624214; DOI: 10.1117/12.2506646 School psychology international. , 2019, Vol.10889, p.16
    • Gritton, H.J., Howe, W.M., Romano, M.F. et al. Unique contributions of parvalbumin and cholinergic interneurons in organizing striatal networks during movement. Nat Neurosci 22, 586–597 (2019).
    • James, Nicholas M; Gritton, Howard J; Kopell, Nancy; Sen, Kamal; Han, Xue ISSN: 0028-3908 , 1873-7064; DOI: 10.1016/j.neuropharm.2018.10.027 Neuropharmacology. , 2019, Vol.144, p.155-171
    • Michael Romano, Mark Bucklin, Howard Gritton, Dev Mehrotra, Robb Kessel, Xue Han, A Teensy microcontroller-based interface for optical imaging camera control during behavioral experiments, Journal of Neuroscience Methods, Volume 320, 2019, Pages 107-115, ISSN 0165-0270, (
    • Piatkevich KD, Bensussen S, Tseng HA, Shroff SN, Lopez-Huerta VG, Park D, Jung EE, Shemesh OA, Straub C, Gritton HJ, Romano MF, Costa E, Sabatini BL, Fu Z, Boyden ES, Han X. Population imaging of neural activity in awake behaving mice. Nature. 2019 Oct;574(7778):413-417. doi: 10.1038/s41586-019-1641-1. Epub 2019 Oct 9. PMID: 31597963; PMCID: PMC6858559.


    • Xiao, S., Tseng, Ha., Gritton, H. et al. Video-rate volumetric neuronal imaging using 3D targeted illumination. Sci Rep 8, 7921 (2018).


    • Wang, Guan et al. “Crucial Roles for SIRT2 and AMPA Receptor Acetylation in Synaptic Plasticity and Memory.” Cell reports. 20.6 (2017): 1335–1347. Web.


    • Jia-Min, Zhuo, Tseng Hua-an, Desai Mitul, Mark E. Bucklin, Ali I. Mohammed, T. M. Robinson Nick, Edward S. Boyden, et al. 2016. "Young Adult Born Neurons Enhance Hippocampal Dependent Performance Via Influences on Bilateral Networks." ELife 5. doi:
    • Mohammed, A., Gritton, H., Tseng, Ha. et al. An integrative approach for analyzing hundreds of neurons in task performing mice using wide-field calcium imaging. Sci Rep 6, 20986 (2016).
    • Sarter, Martin et al. “What Do Phasic Cholinergic Signals Do?” Neurobiology of learning and memory. 130 (2016): 135–141. Web.


    • Ajeesh Koshy Cherian, Howard Gritton, David E. Johnson, Damon Young, Rouba Kozak, Martin Sarter, A systemically-available kynurenine aminotransferase II (KAT II) inhibitor restores nicotine-evoked glutamatergic activity in the cortex of rats, Neuropharmacology, Volume 82, 2014, Pages 41-48, ISSN 0028-3908, (
    • Kopell, Nancy J et al. “Beyond the Connectome: The Dynome.” Neuron. 83.6 (2014): 1319–1328. Web.
    • Kopell, Nancy J; Gritton, Howard J; Whittington, Miles A; Kramer, Mark A ISSN: 0896-6273 , 1097-4199; DOI: 10.1016/j.neuron.2014.08.016 Neuron. , 2014, Vol.83(6), p.1319-1328
    • Lisa A. Briand, Howard Gritton, William M. Howe, Damon A. Young, Martin Sarter, Modulators in concert for cognition: Modulator interactions in the prefrontal cortex, Progress in Neurobiology, Volume 83, Issue 2, 2007, Pages 69-91, ISSN 0301-0082, (
    • Sarter, M., Lustig, C., Howe, W.M., Gritton, H. and Berry, A.S. (2014), Deterministic functions of cortical acetylcholine. Eur J Neurosci, 39: 1912-1920.


    • Gritton HJ, Stasiak AM, Sarter M, Lee TM (2013) Cognitive Performance as a Zeitgeber: Cognitive Oscillators and Cholinergic Modulation of the SCN Entrain Circadian Rhythms. PLoS ONE 8(2): e56206.


    • Gritton, Howard J. 2011. "Bi-Directional Interactions between Cognition and Circadian Rhythms." Order No. 3476482, University of Michigan.


    • Gritton, Howard J et al. “Interactions between Cognition and Circadian Rhythms: Attentional Demands Modify Circadian Entrainment.” Behavioral neuroscience. 123.5 (2009): 937–948. Web.


    • Rose, G., Goller, F., Gritton, H. et al. Species-typical songs in white-crowned sparrows tutored with only phrase pairs. Nature 432, 753–758 (2004).