Biochemistry and Molecular Biology
Penn State Science
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Ying Gu

Ying Gu

Main Content

  • Associate Professor of Biochemistry and Molecular Biology
262 North Frear Laboratory
University Park, PA 16802
Phone: (814) 867-3827


Research Interests

Mechanism of cellulose biosynthesis in arabidopsis

Graduate Programs


Research Summary

Cellulose is the most abundant biopolymer on earth. The great abundance of cellulose places it at the forefront as a primary source of biomass for renewable biofuels and a variety of efforts are underway to improve cellulose degradation. However, little is known about the mechanism by which plant cells make cellulose. Understanding the complex process of cellulose synthesis will be important for optimizing the use of cellulose as a renewable energy source. Cellulose microfibrils are synthesized at the plasma membrane by hexameric protein complexes, also known as cellulose synthase complexes (CSCs). The only known components of CSCs are cellulose synthase (CESA) proteins, first discovered in bacteria in 1990. The principal investigator recently identified a novel plant gene, CSI1, which associates with CESA complexes and are required for normal cellulose biosynthesis. CSI1, as the first non-CESA proteins associated with CSCs, opens up many opportunities. The successful identification of CSI1 prompts us to further explore molecular genetics and biochemical approaches in identification of additional players in CSCs. The cutting-edge live cell imaging will be used to visualize CSCs in living plant cells and to assess individual components’ function in CSCs. Together with biochemical, molecular genetics, and plant genetics approaches, we will pursue the following objectives aimed to unravel the mystery of cellulose biosynthesis in plants: 1) Identification and characterization of novel components in CSCs. 2) Investigate interactions between minimal components in CSCs. 3) Advance our understanding in assembly, delivery, and regulation of CSCs. Together, these studies will substantially increase our knowledge of how plant cells make cellulose and provide unprecedented perspective that aids to increase the efficiency of biomass-based energy production.


Gu figure 1

Figure 1 Imaging of CESA complexes (CSCs). (A) Hexameric CSCs, also known as rosettes, are observed by freeze fracture electron microscopy in algae, moss, and vascular plants. Images are adapted from Giddings et al. (Giddings et al., 1980). (B) CSCs are thought to be composed of 36 subunits of three types in vascular plants, with a diameter about 30 nm. (C) In vivo imaging of CSCs in Arabidopsis. CSI1 is the first non-CESA protein co-localized with CSCs. Bar = 5 μM.



Figure 2-Gu


Figure 2 Hypothetical schematic diagram of the trafficking of CSCs to and from the plasma membrane. CSCs are presumably synthesized in ER and delivered to Golgi for assembly. From Golgi to plasma membrane, delivery may occur directly from the trans-Golgi network (TGN) or through an intermediate compartment such as the MASC/SmaCC. At the plasma membrane, CESA interactive proteins, e.g. CSI1, bridge between CSCs and microtubules and enforce the co-alignment of newly synthesized cellulose microfibrils and cortical microtubules. Adapted from Lei et al., 2012.

Representative Publications

Selected Publications


• Lei L, Singh A, Bashline L, Li S, Yingling YG, Gu Y (2015) The Arabidopsis cellulose synthase interactive 1 is required for a fast recycling of cellulose synthase complex to the plasma membrane. Plant Cell pii: tpc.15.00442.
• 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, 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
• 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 mediate cellular response to the anticancer PAD4 inhibitor YW3-56. Mol. Cancer Ther. 14:877-888. PMID: 25612620
• 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.
• 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, 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
• Ye X, Lei L, Stork J, Brabham C, Strickland J, Ladak A, Gu Y, Debolt S (2014) Microbial natural product-based bioprospecting: identification of acetobixan as an inhibitor of plant cellulose biosynthesis. PLoS ONE DOI: 10.1371/journal.pone.0095245
• 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, Li S, Gu Y (2014) Trafficking of the cellulose synthase complex in higher plants. Ann. Bot. doi: 10.1093/aob/mcu040
• Bashline L, Lei L, Li S, Gu Y (2014) Cell wall, cytoskeleton, and cell expansion in higher plants.  Mol. Plant 7(4): 586-600
• Li S, Bashline L, Lei L, Gu Y (2014) Cellulose biosynthesis and its regulation. The Arabidopsis Book 11:e0169. doi: 10.1199/tab.0169
• 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 ?2, 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 Z (2012) AtPRK2 promotes ROP1 activation via RopGEFs in the control of polarized pollen tube growth. Mol. Plant 6(4): 1187-1201*Joint first authors
• Li S and Gu Y (2012) Cellulose biosynthesis in higher plants and the role of the cytoskeleton. eLS 8(3):e23179
• 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
• 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
• 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
• 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
• Lei L, Li S, Gu Y (2012) Cellulose synthase complexes: composition and regulation. Front. Plant Physiol. 3: 75 doi: 10.3389/fpls.2012.00075
• 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.
• Bashline L, Du J, Gu Y (2011) The trafficking and behavior of cellulose synthase and a glimpse of potential cellulose synthesis regulators. Front. Biol. 6(5): 377-383
• Gu Y*, Somerville C (2010) Cellulose synthase interacting protein: a new factor in cellulose synthesis. Plant Signal. Behav. 5(12): 1571-1574
• *Corresponding author
• Gu Y, Kaplinsky N, Bringmann M, Cobb A, Carroll A, Sampathkumar A, Baskin TI, Persson S, Somerville C (2010) Identification of a cellulose synthase-associated protein required for cellulose biosynthesis. Proc. Natl. Acad. Sci. 107(29): 12866-12871
• Gu Y, Deng ZP, Paredez AR, Debolt S, Wang ZY, Somerville C (2008) Prefoldin6 is required for normal microtubule dynamics and organization in Arabidopsis. Proc. Natl. Acad. Sci. 105(46): 18064-18069
• Li S, Gu Y, Yan A, Lord E, Yang ZB (2008) RIP1 (ROP Interactive Partner 1)/ICR1 marks pollen germination sites and may act in the ROP1 pathway in the control of polarized pollen growth. Mol. Plant 6:1021-1035
• Jeon BW, Hwang JU, Hwang YK, Song WY, Fu Y, Gu Y, Bao F, Cho D, Kwak JM, Yang ZB, Lee Y (2008) The Arabidopsis small G protein ROP2 is activated by light in guard cells and inhibits light-induced stomatal opening. Plant Cell 20: 75-87
• Hwang JU, Gu Y, Lee YJ, Yang ZB (2006) A Oscillatory ROP GTPase activation leads the oscillatory polarized growth of pollen tubes. Mol. Biol. Cell 16: 5385-5399
• Gu Y, Li SD, Lord EM, Yang ZB (2006) Members of a novel class of Arabidopsis Rho guanine nucleotide exchange factors control Rho GTPase-dependent polar growth. Plant Cell 18: 366-381
• Fu Y*, Gu Y*, Zheng ZL, Wasteneys G, Yang ZB (2005) Arabidopsis interdigitating cell growth requires two antagonistic pathways with opposing action on cell morphogenesis. Cell 120: 687-700 * Joint first authors
• Gu Y, Fu Y, Dowd P, Li SD, Vernoud V, Gilroy S, Yang ZB (2005) A Rho-family GTPase controls actin dynamics and tip growth via two counteracting downstream pathways in pollen tubes. J. Cell Biol. 169:127-138
• Gu Y, Wang ZH, Yang ZB (2004) ROP/RAC GTPase: an old new master regulator for plant signaling. Curr. Opin. Plant Biol. 7: 527-536
• Park J, Gu Y, Lee Y, Yang ZB, Lee Y (2004) Phosphatidic acid induces leaf cell death in Arabidopsis by activating the Rho-related small G protein GTPase-mediated pathway of reactive oxygen species generation. Plant Physiol. 134: 129-136
• Gu Y, Vernoud V, Fu Y, Yang ZB (2003) ROP GTPase regulation of pollen tube growth through the dynamics of tip-localized F-actin. J Exp. Bot. 54: 93-101
• Wu G*, Gu Y*, Li SD, Yang ZB (2001) A genome-wide analysis of Arabidopsis Rop-interactive CRIB motif-containing proteins that act as Rop GTPase targets. Plant Cell 13: 2841-2856 * Joint first authors