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

  • 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): 11383-11353
  • Wang C, Hu T, Yan X, Meng T, Wang Y, Wang Q, Zhang X, Gu Y, Sánchez-Rodríguez C, Gadeyne A, Lin J, Persson S, Van Damme D, Li C, Bednarek SY, Pan J (2016) Differential regulation of Clathrin and its adaptor proteins during their membrane recruitment in Arabidopsis. Plant Physiol. 171(1): 215-229
  • 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
  • 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
  • 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, 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.
  • 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
  • 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
  • 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