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Ying Gu

Associate Department Head for Graduate Education and Professor of Biochemistry and Molecular Biology
Dr. Ying Gu

About Me

Dr. Gu received her PhD in Plant Genetics at University of California-Riverside (supervisor: Dr. Zhenbiao Yang). For her thesis work, she made important contributions at the molecular level to the understanding of polar growth (pollen tube) and diffuse growth (pavement cells). She worked as a postdoctoral fellow at Stanford University (supervisor: Dr. Chris Somerville) and as an associate specialist at University of California Berkeley when Dr. Somerville became the Director of Energy Biosciences Institute. Dr. Gu started her own lab at Penn State in 2010. Her group is part of the “Center for Lignocellulose Structure and Formation”, an Energy Frontier Research Center.


Program or Department Affiliations

BMMB Graduate Program Molecular, Cellular, and Integrative Biosciences Plant Biology


Center for Lignocellulose Structure and Formation Center for Cellular Dynamics

Research Interest

Molecular mechanisms of cellulose biosynthesis in plants


Research Summary

Gu Lab uses biochemical, molecular genetics, and spectroscopic approaches to decipher mechanisms of cellulose biosynthesis. Cellulose is the most abundant biopolymer on earth. Cellulose is a major source of raw materials for paper, textiles, and an abundant source for sustainable and environment-friendly cellulosic biofuels. A long-term goal of our research is to gain a deeper understanding of the regulation of cellulose biosynthesis so that the fundamental knowledge can be transferred for designing new cellulosic materials with diverse economic applications.


Honors, Awards and Professional Leadership

  • 2020 Daniel R. Tershak Memorial Teaching Award
  • 2017-present Editorial Board Member, Plant Direct 
  • 2011-present Editorial Board Member, Frontiers in Plant Physiology  
  • 2011-present Editorial Board Member, Journal of Glycobiology


Selected Publications

  • Allen H, Zhu X, Li S, Gu Y (2024) The TRAPPIII subunit, Trs85, has a dual role in the trafficking of cellulose synthase complexes in Arabidopsis. Plant Journal
  • Allen H, Davis B, Patel J, Gu Y (2024) Dot Scanner: Open-source software for quantitative live-cell imaging in planta. Plant Journal
  • Rongpipi S, Barnes WJ, Siemianowski O, Ye D, Del Mundo JT, Duncombe S, Xin X, Zhu C, Toney MF, Gu Y Anderson CT, Gomez ED, Gomez EW (2024) Matrix Polysaccharides affect preferred orientation of cellulose crystals in primary cell walls. Cellulose  31, 1397-1415.
  • Lee J, Choi J, Feng L, Yu J, Zheng Y, Zhang Q, Lin YT, Sah S, Gu Y, Zhang S, Cosgrove DJ, Kim SH (2023) Regiospecific cellulose orientation and anisotropic mechanical property in plant cell walls. Biomacromolecules 24, 4759-4770
  • Xin X*, Wei D*, Lei L, Zheng H, Wallace IS, Li S, Gu Y (2023) CALCIUM-DEPENDENT PROTEIN KINASE32 regulates cellulose biosynthesis through post-translational modification of cellulose synthase. New Phytologist 239, 2212-2224. *co-first author
  • Murshed M, Wei D, Gu Y*, Wang J* (2023) Simulation of microtubule-cytoplasm interaction revealed the importance of fluid dynamics in determining the organization of microtubules. Plant Direct 7, e505. *Corresponding author
  • Gu Y*, Rasmussen C* (2022) Cell biology of primary cell wall synthesis in plants. Plant Cell 34, 103-128. *Corresponding author
  • Allen H, Wei D, Gu Y, Li S (2021) A historical perspective on the regulation of cellulose biosynthesis. Carbohydrate Polymers 252, 117022-107039.
  • Zhu X, Tellier F, Gu Y*, Li S* (2020) Disruption of very-long-chain-fatty acid synthesis has an impact on the dynamics of cellulose synthase in Arabidopsis thaliana. Plants 9, 1599-1618. *Corresponding author
  • Makarem M, Nishiyama Y, Xin X, Durachko D, Gu Y, Cosgrove D, Seong K (2020) Distinguishing mesoscale polar order (Unidirectional vs. Bidirectional) of cellulose microfibrils in plant cell walls using sum frequency generation spectroscopy. J. Phys. Chem. 124, 37:8071-8081.
  • Xin X, Lei L, Zheng Y, Zhang T, Pingali SV, O’Neil H, Cosgrove DJ, Li S, Gu Y (2020) CELLUOSE SYNTHASE INTERACTIVE1 is required for the crossed-polylamellate wall architecture of Arabidopsis outer epidermal cell walls. J. Exp. Bot. 71:2982-2994.
  • Zhu X, Xin X, Gu Y (2019) Cellulose and hemicellulose synthesis and their regulation in plant cells. Book chapter, Extracellular sugar-based biopolymer matrices. DOI 10.1007/978-3-030-12919-4, Ephraim Cohen and Hans Merzendorfer, ed. Springer Nature Switzerland AG.
  • Phyo P, Gu Y, Hong M (2019) Impact of acidic pH on plant cell wall polysaccharide structure and dynamics: insights into the mechanism of acid growth in plants from solid-state NMR. Cellulose 26:291-304.
  • Zhu X, Li S, Pan S, Xin X, Gu Y (2018) CSI1, PATROL1 and exocyst complex cooperate in delivery of cellulose synthase complexes to the plasma membrane. Proc. Natl. Acad. Sci. USA 115:E3578-3587.
  • Li S, Bashline L, Zheng Y, Xin X, Huang S, Kong Z, Kim SH, Cosgrove D, Gu Y (2016) Cellulose synthase complexes act in a concerted fashion to synthesize highly aggregated cellulose in secondary cell walls of plants. Proc. Natl. Acad. Sci. USA 113(40): 11348-11353
  • Lei L, Singh A, Bashline L, Li S, Yingling YG, Gu Y (2015) Cellulose synthase interactive 1 is required for a fast recycling of cellulose synthase complex to the plasma membrane in Arabidopsis. Plant Cell 27(10): 2926-2940
  • Bashline L, Li S, Zhu X, Gu Y (2015) The TWD40-2 protein and the AP2 complex cooperate in the clathrin-mediated endocytosis of cellulose synthase to regulate cellulose biosynthesis. Proc. Natl. Acad. Sci. USA 112(41): 12870-12875
  • Li S, Lei L, Yingling YG, Gu Y (2015) Microtubules and cellulose biosynthesis: the emergence of new players. Curr. Opin. Plant Biol. 28: 76-82
  • Bashline L, Gu Y (2015) Using the Split-Ubiquitin yeast two-hybrid systems to test protein-protein interactions of transmembrane proteins. In: Plant Cell Expansion: Methods and Protocols. Method. Mol. Biol. vol. 1242, DOI 10.1007/978-1-4939-1902-4_13, Jose M. Estevez, ed. Springer Science, New York.
  • Wang S, Chen XA, Hu J, Jiang JK, Li Y, Chan-Salis KY, Gu Y, Chen G, Thomas C, Pugh BF, Wang Y (2015) ATF4 gene network mediates cellular response to the anticancer PAD4 inhibitor YW3-56 in triple negative breast cancer cells. Mol. Cancer Ther. 14:877-888. PMID: 25612620
  • Brabham C*, Lei L*, Stork J, Barrett M, Gu Y, Debolt S (2014) Indaziflam herbicidal action: a potent cellulose biosynthesis inhibitor. *Joint first authors Plant Physiol. 166(3): 1177–1185
  • Lei L, Li S, Bashline L, Gu Y (2014) Dissecting the molecular mechanism underlying intimate relationship between cellulose microfibrils and cortical microtubules. Front. Plant Sci. doi: 10.3389/fpls.2014.00090
  • Bashline L, Lei L, Li S, Gu Y (2014) Cell wall, cytoskeleton, and cell expansion in higher plants. Mol. Plant 7(4): 586-600
  • Lei L, Zhang T, Strasser R, Lee CM, Gonneau M, Mach L, Vernhettes S, Kim SH, Cosgrove D, Li S, Gu Y (2014) The jiaoyao1 mutant is an allele of korrigan that abolishes endoglucanase activity and affects the organization of both cellulose microfibrils and microtubules in Arabidopsis. Plant Cell 26(6): 2601-2616
  • Li S, Bashline L, Lei L, Gu Y (2014) Cellulose biosynthesis and its regulation. The Arabidopsis Book 11:e0169. doi: 10.1199/tab.0169
  • Ye X, Lei L, Stork J, Brabham C, Strickland J, Ladak A, Gu Y, Debolt S (2014) Acetobixan, an inhibitor of cellulose synthesis identified by microbial bioprospecting. PLoS ONE DOI: 10.1371/journal.pone.0095245
  • Bashline L, Li S, Gu Y (2014) Trafficking of the cellulose synthase complex in higher plants. Ann. Bot. doi: 10.1093/aob/mcu040
  • Lei L, Li S, Juan Du, Bashline L, Gu Y (2013) Cellulose synthase interactive 3 regulates cellulose biosynthesis in both microtubule-dependent and microtubule-independent manner. Plant Cell 25(12): 4912-4923
  • Bashline L, Li S, Anderson CT, Lei L, Gu Y (2013) The endocytosis of cellulose synthase in Arabidopsis is dependent on m2, a clathrin mediated endocytosis adaptin. Plant Physiol. 163(1): 150-160
  • Li S, Lei L, Gu Y (2013) Functional analysis of complexes with mixed primary and secondary cellulose synthases. Plant Signal. Behav. 8(3). pii: e23179
  • Chang F*, Gu Y*, Ma H, Yang ZB (2013) AtPRK2 promotes ROP1 activation via RopGEFs in the control of polarized pollen tube growth. Mol. Plant 6(4): 1187-1201*Joint first authors
  • Lei L, Li S, Gu Y (2012) Cellulose synthase complexes: composition and regulation. Front. Plant Sci. 3: 75 doi: 10.3389/fpls.2012.00075
  • Carroll A, Mansoori N, Li S, Lei L, Vernhettes S, Visser R, Somerville C, Gu Y, Trindade L (2012) Complexes with mixed primary and secondary cellulose synthases are functional in planta. Plant Physiol.  160(2): 726-737
  • Li S and Gu Y (2012) Cellulose biosynthesis in higher plants and the role of the cytoskeleton. eLS 8(3):e23179
  • Wang Y, Li P, Wang S, Hu J, Chen XA, Wu J, Fisher M, Oshaben K, Zhao N, Gu Y, Chen G, Wang Y. (2012) Anticancer PAD inhibitors regulate autophagy and the mammalian target of rapamycin complex 1 activity. J. Biochem. Chem. 287(31): 25941-25953
  • Baskin T and Gu Y (2012) Making parallel lines meet: Transferring information from microtubules to extra-cellular matrix. Cell Adh. Migr. 6(5): 1-5
  • Lei L, Li S, Gu Y (2012) Cellulose synthase interactive protein 1 (CSI1) mediates the intimate relationship between cellulose microfibrils and cortical microtubules. Plant Signal. Behav. 7(7): 714-718
  • Li S, Lei L, Somerville C, Gu Y (2012) Cellulose synthase interactive protein 1 (CSI1) links microtubules and cellulose synthase complexes. Proc. Natl. Acad. Sci. 109 (1) 185-190.