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  Location: Home >> Faculty >> Center for Genome Biology
  Center for Genome Biology


Yihua Zhou


Dr. Yihua Zhou, Principle Investigator,
B.S. 1985-1989, Department of Biology, Beijing Normal University
M.S. 1989-1992, Department of Biology, Beijing Normal University
Ph. D. 1996-1999, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences.
Postdoctoral research experience 2001-2002, Department of Forestry, Michigan Technological University.
 
Tel: 64807605 (O); 64807628 (L)
Email: yhzhou@genetics.ac.cn.


ESEARCH FOCUS
 
Functional characterization of genes involved in secondary wall biosynthesis to improve lodging resistance in crops
Lodging resistance is an important trait that is highly associated with crop yield. Many genes have been found responsible for this trait, in which genes involved in secondary wall biosynthesis make a great contribution, as physicochemical property of secondary wall determines the mechanical strength of plants. My laboratory is mainly interested in molecular mechanism of secondary wall formation in rice developing internodes, focusing on functional characterization of genes involved in secondary wall biosynthesis and remodeling. Therefore, we systematically screen rice brittle culm mutants (bc) and functionally characterize the genes related to mechanical properties and cell wall biosynthesis. Till now, eight genes: BC1 (Liu et al., 2013), BC15 (Wu et al., 2012), BC13 (Song et al., 2012), BC14 (Zhang et al., 2011), BC3 (Xiong et al., 2010), BC12 (Zhang et al., 2010), and BC11 (Zhang et al., 2009) have been identified, which have drawn the major framework of cell wall biosynthesis and allow us stride a step in controlling mechanical strength of rice plants via molecular breeding.
 
On the other hand, plant cell wall is a natural nanoscale network structure. How plants organize the cell wall polymers determines mechanical properties of the cell wall. As the developments of nuclear magnetic resonance spectroscopy (NMR) and atomic force microscopy (AFM), we and our colleagues found xylan an important cross-link polysaccharide that can bind cellulose and lignin via different conformations. Acetylation profile on xylan is a crucial manner to regulate the conformation of xylan. In recent years, we already identified a xylan backbone acetyltransferase OsTBL1/2 (Gao et al., 2017), a backbone esterase BS1 (Zhang et al., 2017), and a sidechain esterase (Gao et al., 2019). Functional characterization of more proteins involved in the xylan acetylation profile control may provide a tool for precise improvement of agronomic traits, like lodge resistance.
 
Hemicellulose biosynthesis and modification
Xylan and mixed-linkage glucan (MLG) are major hemicelluloses present in rice cell wall. Both polymers have β-1,4-xylosyl or glucosyl backbone, respectively. Xylan backbone is further O-acetylated, which determines xylan physicochemical properties. However, the molecular mechanisms about how both hemicellulosic polymers are synthesized in plants and by which components control the acetyl modification on xylan are poorly understood. Our laboratory focuses on identifying the critical genes required for xylan synthesis and understanding their biological functions in rice. We have found several glycosyltransferases functioning in xylan synthesis through genetic characterization. We recently reported that rice Trichome Birefringence-Like1/2 (OsTBL1/2) and Brittle Sheath1 (BS1) function as xylan acetyltransferase and acetylesterase, respectively (Gao et al., 2017; Zhang et al., 2017), indicating that xylan acetylation control involves acetyl-addition and -removal processes. These studies outline a bidirectional regulation mechanism for how plants modulate the acetyl-ester profile on xylan and shed light on understanding their corresponding significance in plant growth and economic values.
 
For the synthesis of MLG, we have identified some natural rice varieties that possess high level of MLG. We propose to use forward genetic approach to clone the candidate gene, aiming to clarify the new mechanisms for MLG biosynthesis and its function.
 
Cell wall biology and plant growth
Cell wall is a basic structure of plant cells. Physicochemical properties of cell wall highly fit with their roles of cells across different growth and development stages. Therefore, plant cell walls play many important roles during plant growth and development. Deficiency in cell wall biosynthesis often exhibits abnormalities in plant growth. However, the underlying mechanisms of how plants perceive signals to initiate cell wall formation and remodeling and how the signaling pathways transmit are largely unknown. In the recent years, our laboratory is focusing on characterizing cell wall-related genes required for plant growth and identifying the signaling transduction pathways that regulate cell wall formation and plant growth. For example, ILA1 (Ning et al., 2011) and IIP4 (Zhang et al., 2018) mediated ILA1-IIP4-NAC function in mechanical tissue formation. Gibberellin (GA) is an important phytohormone. Our recent work found that GA can regulate secondary wall cellulose biosynthesis. A signal transduction pathway that contains SLR1-NAC29/31-MYB61-secondary wall CESAs has been identified, which is conserved in many plant species (Huang et al., 2015). KNAT7 (Wang et al., 2019) coordinates cell expansion and strengthening.
Those findings forward our knowledge about the relationship between plant growth and cell wall biosynthesis.
 
RESEARCH TEAM
Assistant Professor: Dr. Baocai Zhang
Assistants:        Dr. Lanjun Zhang              
Ms. Xiangling Liu
PhD students: Yihong Gao, Hanlei Yang, Chenxu Gao, Jianing Ma, Zhao Wen, Zhuolin Liu, Yan Wang, Jiaxi Li
MS students: Jiaxin Ma
Guest student/Postdoc: Hang Wang, Shaoxue Chao, Zuopeng Xu, Rui Xu
 

RECENT PUBLICATIONS
2022:
Zhou Y and Zhang B. (2022) Unprecedented polysaccharide nanostructures sustain vessel wall patterning and robustness. Nat Plants 8, 330–331(Research Briefing).
 
Wang H, Yang H, Wen Z, Gao C, Gao Y, Tian Y, Xu Z, Liu X, Persson S, Zhang B*, Zhou Y*. (2022) Xylan-based nanocompartments orchestrate plant vessel wall patterning. Nat Plants 8, 295–306.
 
Wang F, Cheng Z, Wang J, Zhang F, Zhang B, Luo S, Lei C, Pan T, Wang Y, Zhu Y, Wang M, Chen W, Lin Q, Zhu S, Zhou Y, et al. (2022) Rice STOMATAL CYTOKIESIS DEFECTIVE2 regulates of cell expansion by affecting vesicular trafficking in rice. Plant Physiol. 189, 567–584.
 
Qiang Z, Sun H, Ge F, Li W, Li C, Wang S, Zhang B, Zhu L, Zhang S, Wang X, Lai J, Qin F, Zhou Y, Fu Y. (2022) The transcription factor ZmMYB69 represses lignin biosynthesis by activating ZmMYB31/42 expression in maize. Plant Physiol. In press doi.org/10.1093/plphys/kiac233
 
2021:
 
Zhang B, Gao Y, Zhang L, Zhou Y* (2021). The plant cell wall: biosynthesis, construction, and functions. J. Integr. Plant Biol. 63: 251–272
 
2020:
 
Gao Y, Xu Z, Zhang L, Li S, Wang S, Yang H, Liu X, Zeng D, Liu Q, Qian Q, Zhang B*, Zhou Y*(2020). GRF4-regulated MYB61 improves nitrogen utilization and biomass production in rice. Nat. Commun. 11:5219.
 
 
Xue J#, Zhang B#, Zhan H#, Lv Y, Jia X, Wang T, Yang N, Lou Y, Zhang Z, Hu W, Gui J, Cao J, Xu P, Zhou Y, Hu J, Li L*, Yang Z*. (2020). Phenylpropanoid derivatives are essential components of sporopollenin in vascular plants. Mol. Plant 13: 1644–1653.
 
Zhao M, Tang S, Zhang H, He M, Liu J, Zhi H, Sui Y, Liu X, Jia G, Zhao Z, Yan J, Zhang B. Zhou Y, Chu J, Wang X, Zhao B, Tang W, Li J, Wu C, Liu X*, Dao X*. (2020). DROOPY LEAF1 controls leaf architecture by orchestrating early brassinosteroid signaling. Proc. Natl. Acad. Sci. USA 117, 21766–21774.
 
Zhao X, Ebert B, Zhang B, Liu H, Zhang Y, Zeng W, Rautengarten C, Li H, Chen X, Bacic A, Wang G, Men S, Zhou Y, Heazlewood J, Wu A*. (2020). UDP‐Api/UDP‐Xyl synthases affect plant development by controlling the content of UDP‐Api to regulate the RG‐II‐borate complex. Plant J. 104, 252–267.
 
2019
 
Zhang L, Gao C, Mentink-Vigier F, Tang L, Zhang D, Wang S, Cao S, Xu Z, Liu X, Wang T, Zhou Y*, and Zhang B*. (2019) Arabinosyl deacetylase modulates the arabinoxylan acetylation profile and secondary wall formation. Plant Cell 31, 1113–1126.
 
Wang S, Yang H, Mei J, Liu X, Wen Z, Zhang L, Xu Z, Zhang B*, and Zhou Y*. (2019) Rice homeobox protein KNAT7 integrates the pathways regulating cell expansion and wall stiffness. Plant Physiol. 181, 669–682.
 
Zhang L, Zhang B*, and Zhou Y*. (2019). Cell wall compositional analysis of rice culms. Bio-protocol 9: e3398.
 
2018:
 
Zhang Z, Zhang B, Chen Z, Zhang D, Zhang H, Wang H, Zhang Y, Cai D, Liu J, Xiao S, Huo Y, Liu Jie, Zhang L, Wang M, Liu X, Xue Y*, Zhao L*, Zhou Y*, and Chen H*. (2018) A PECTIN METHYLESTERASE at the Maize Ga1 locus confers male function in unilateral cross-incompatibility. Nat. Commun. 9, 3678
 
Zhang D, Xu Z, Cao S, Chen K, Li S, Liu X, Gao C, Zhang B*, and Zhou Y*. (2018) An uncanonical CCCH-tandem zinc finger protein represses secondary wall synthesis and controls mechanical strength in rice. Mol. Plant 11, 163–174.
 
Sanchez-Rodriguez C.*, Shi Y., Kesten, C., Zhang, D., Sancho-Andres, G., Ivakov, A., Lampugnani, E., Sklodowski, K., Fujimoto, M., Nakano, A., Bacic, A., Wallace, I., Ueda, T., Damme, D., Zhou, Y., and Persson S. (2018) The cellulose synthases are cargo of the TPLATE adaptor complex. Mol. Plant 11, 346–349.
 
Zhang L-M, Leng C-Y, Luo H, Wu X-Y, Liu Z-Q, Zhang Y-M, Zhang H, Xia Y, Shang L, Liu C-M, Hao D-Y, Zhou Y, Chu C, Cai H-W, and Jing H-C*. (2018) Sweet sorghum originated through selection of dry, a plant-specifific NAC transcription factor gene. Plant Cell 30, 2286–2307.
 
Xu, J., Zhang, B., Lu, X., Zhou, Y., Fang, J., Li Y., and Zhang, S*. (2018) Nanoscale observation of microfibril swelling and dissolution in ionic liquids. ACS Sustainable Chem. Eng. 6, 909–917.
 
Zhang C, Shen Y, Tang D, Shi W, Zhang D, Du G, Zhou Y, Liang G, Li Y, and Cheng Z*. (2018) The zinc finger protein DCM1 is required for male meiotic cytokinesis by preserving callose in rice. PLoS Genet. (DOI:10.1371/journal.pgen.1007769 )
 
2017:
 
Zhang B, Zhang L, Li F, Zhang D, Liu X, Wang H, Xu Z, Chu C*, and Zhou Y*. (2017) Control of secondary cell wall patterning involves xylan deacetylation by a GDSL esterase. Nat. Plants 3,17017.
 
Gao Y, He C, Zhang D, Liu X, Xu Z, Tian Y, Liu X, Zang S, Pauly M, Zhou Y*, and Zhang B*. (2017) Two trichome birefringence-like proteins mediate xylan acetylation, which is essential for leaf blight resistance in rice. Plant Physiol. 173, 470–481.
 
Zhang B and Zhou Y*. (2017) Carbohydrate composition analysis in xylem. de Lucus M. and Etchells JP, eds. Humana Press. New York. Methods Mol. Biol. 1544: 213–222.
 
Xu Z#, Li S#, Zhang C, Zhang B, Zhu K, Zhou Y*, and Liu Q*. (2017) Genetic connection between cell-wall composition and grain yield via parallel QTL analysis in indica and japonica subspecies. Sci. Rep. 7: 12561.
 
Schneider R, Tang L, Lampugnani E R, Barkwill S, Lathe R, Zhang Y, McFarlane HE,  Pesquet E, Niittyla T, Mansfield SD, Zhou Y, and Persson S. (2017) Two complementary mechanisms underpin cell wall patterning during xylem vessel development. Plant Cell 29: 2433–2449.
 
Qi J, Wu B, Feng S, Lü S, Guan C, Zhang X, Qiu D, Hu Y, Zhou Y, Li C, Long M*, and Jiao Y* (2017) Mechanical regulation of organ asymmetry in leaves. Nat. Plants 3: 724–733.
 
Zhou T, Hua Y, Zhang B, Zhang X, Zhou Y, Shi L, and Xu F*. (2017) Low-boron tolerance strategies involving pectin-mediated cell wall mechanical properties in Brassica napus. Plant Cell Physiol. 58: 1991–2005.
 
2016:
 
Ma J, Cheng Z, Chen J, Shen J, Zhang B, Ren Y, Ding Y, Zhou Y, Zhang H, Zhou K, Wang J, Lei C, Zhang X, Guo X, Gao H, Bao Y, Wan J*. (2016) Phosphatidylserine synthase controls cell elongation especially in the uppermost internode in rice by regulation of exocytosis. PLoS ONE 11, e0153119.
 
2015:
 
Huang D, Wang S, Zhang B, Shang-Guan K, Shi Y, Zhang D, Liu X, Wu K, Xu Z, Fu X, and Zhou Y. (2015) A Gibberellin-midiated DELLA-NAC signaling cascade regulates cellulose synthesis in rice. Plant Cell 27: 1681-1696.
 
张保才, 周奕华*. (2015). 植物细胞壁形成机制的新进展.中国科学, 45: 544-556.
 
Liu S., Lan J., Zhou B., Qin Y., Zhou Y., Xiao X., Yang J., Gou J., Qi J., Huang Y., Tang C. (2015) HbNIN2, a cytosolic A/N-invertase, is responsible for sucrose catabolism in rubber-producing laticifers of Hevea brasiliensis (para rubber tree). New Phytol. 206: 709–725.
 
Wang X., Jing Y, Zhang B, Zhou Y, and Lin, R*. (2015). Glycosyltransferase-like protein ABI8/ELD1/KOB1 promotes Arabidopsis hypocotyl elongation through regulating cellulose biosynthesis. Plant Cell Environ. 38: 411–422.
 
2014:
 
Shi Y, Liu X, Li R, Gao Y, Xu Z, Zhang B*, and Zhou Y. (2014) Retention of OsNMD3 in the cytoplasm disturbs protein synthesis efficiency and affects plant development in rice. J. Exp. Bot. 65: 3055–3069
 
Wang C, Li S, Ng S, Zhang B, Zhou Y, Whelan J, Wu P, and Shou H*. (2014). Mutation in xyloglucan 6-xylosytransferase results in abnormal root hair development in Oryza sativa. J. Exp. Bot. 65, 4149–4157
 
Zhu X, Sun Y, Zhang B, Mansoori N, Wan J, Liu Y, Wang Z, Shi YZ, Zhou Y, and Zheng S*. (2014). TRICHOME BIREFRINGENCE-LIKE27 affects aluminum sensitivity by modulating the O-acetylation of xyloglucan and aluminum-binding capacity in Arabidopsis. Plant Physiol. 166, 181–189
 
2013:
 
Liu L, Shang-Guan K, Zhang B, Liu X, Yan M, Zhang L, Shi Y, Zhang M, Qian Q, Li J, and Zhou Y. (2013) Brittle Culm1, a COBRA-like protein, functions in cellulose assembly through binding cellulose microfibrils. PLoS Genet. 9 (8): e1003704
 
Song X, Liu L, Jiang Y, Zhang B, Gao Y, Liu X, Lin Q, Ling H, and Zhou Y. (2013) Disruption of secondary wall cellulose biosynthesis alters cadmium translocation and tolerance in rice plants. Mol. Plant 6:768–780.
 
Kang J, Zhang H, Sun T, Shi Y, Wang J, Zhang B, Wang Z, Zhou Y, and Gu H. (2013) Natural variation of C-repeat-binding factor (CBFs) genes is a major cause of divergence in freezing tolerance among a group of Arabidopsis thaliana populations along the Yangtze River in China. New Phytol. 199: 1069–1080.
 
Zou H, Zhang Y, Wei W, Chen H, Song Q, Liu Y, Zhao M, Wang F, Zhang B, Lin Q, Zhang W, Ma B, Zhou Y, Zhang J, and Chen S. (2013) The transcription factor AtDOF4.2 regulates shoot branching and seed coat formation in Arabidopsis. Biochem J. 449: 373–388.
 
2012:
 
Wu B, Zhang B, Dai Y, Zhang L, Shang-Guan K, Peng Y, Zhou Y*, and Zhu Z*. (2012) Brittle Culm15 encodes a membrane-associated chitinase-like protein required for cellulose biosynthesis in rice. Plant Physiol. 159: 1440–1452 (*Corresponding author)
 
Zhang S-J, Song X-Q, Yu B, Zhang B-C, Sun C-Q, Knox J. P, and Zhou Y-H. (2012) Identification of quantitative trait loci affecting hemicellulose characteristics based on cell wall composition in a wild and cultivated rice species. Mol. Plant 5: 162–175.
 
Zhu X., Shi Y., Lei G., Fry S., Zhang B., Zhou Y., Braam J., Jiang T., Xu X., Mao C., Pan Y., Yang J., Wu P., and Zheng S. (2012) XTH31, encoding an in vitro XEH/XET-active enzyme, regulates aluminum sensitivity by modulating in vivo XET action, cell wall xylolucan content, and aluminum binding capacity in Arabidopsis. Plant Cell 24: 4731–4747.
 
Bao C, Wang J, Zhang R, Zhang B, Zhang H, Zhou Y, and Huang S. (2012) Arabidipsis VILLIN2 and VILLIN3 act redundantly in sclerenchyma development via bundling of actin filaments. Plant J. 71: 962–975
 
Chen M-Q, Zhang A-H, Zhang Q, Zhang B-C, Nan J, Li X, Liu N, Qu H, Lu C-M, Sudmorgen, Zhou Y-H, Xu Z-H, and Bai S-N. (2012) Arabidopsis NMD3 is required for nuclear export of 60S ribosomal subunits and affects secondary cell wall thickening. PLoS ONE 7(4): e35904
 
2011:
Ning J, Zhang B, Wang N, Zhou Y*, and Xiong L*. (2011) Increased leaf angle1, a Raf-Like MAPKKK that interacts with a nuclear protein family, regulates mechanical tissue formation in the lamina joint of rice. Plant Cell 23: 4334–4347 (*Corresponding author)
 
Zhang B, Liu X, Qian Q, Liu L, Dong G, Xiong G, Zeng D, and Zhou Y. (2011) A Golgi nucleotide sugar transporter modulates cell wall biosynthesis and plant growth in rice. Proc. Natl. Acad. Sci. USA. 108: 5110–5115.
 
Song X., Zhang B., and Zhou Y. (2011) Golgi-localized UDP-glucose transporter is required for cell wall integrity in rice. Plant Signaling & Behavior, 6: 1097–1100.
 
Zhang B and Zhou Y. (2011) Study on rice brittleness mutants: A way to open the ‘black box’ of monocot cell wall biosynthesis. J. Integr. Plant Biol. 53: 136–142.
 
Liu X-L, Liu L, Niu Q-K, Xia C, Yang K-Z, Li R, Chen L-Q, Zhang X-Q, Zhou Y*, and Ye D*. (2011) MALE GAMETOPHYTE DEFECTIVE 4 encodes a rhamnogalacturonan II xylosyltransferase and is important for growth of pollen tubes and roots in Arabidopsis. Plant J. 65: 647–660 (*Corresponding author)
 
Lei M, Liu Y, Zhang B, Zhao Y, Wang X, Zhou Y, Raghothama K G, and Liu D. (2011) Genetic and genomic evidence that sucrose is a global regulator of plant responses to phosphate starvation in Arabidopsis. Plant Physiol. 156: 1116–1130.
 
Wang W, Wang Li, Chen C, Xiong G, Tan X-Y, Yang K-Z, Wang Z-C, Zhou Y, Ye D, and Chen L-Q. (2011) Arabidopsis CSLD1 and CSLD4 are required for cellulose deposition and normal growth of pollen tubes. J. Exp. Bot. 62: 5161–5177.
 
Wang Z-P, Xie Z-Q, Zhang B, Hou L-Y, Zhou Y, Li L-H, and Han X-G. (2011) Aerobic and anaerobic nonmicrobial methane emissions from plant material. Environ. Sci. Technol. 45: 9531–9537.
 
2010:
 
Xiong G, Li R, Qian Q, Song X, Liu X, Yu Y, Zeng D, Wan J, Li J and Zhou Y. (2010) The rice dynamin-related protein DRP2B mediates membrane trafficking and thereby plays a critical role in secondary cell wall cellulose biosynthesis. Plant J. 64: 56–70.
 
Zhang M, Zhang B, Qian Q, Yu Y, Li R, Zhang J, Liu X, Zeng D, Li J and Zhou Y. (2010) Brittle Culm12, a dual-targeting Kinesin-4 protein, controls cell cycle progression and wall properties in rice. Plant J. 63: 312–328.
 
Li R, Xiong G, and Zhou Y. (2010) Membrane trafficking mediated by OsDRP2B is specific for cellulose biosynthesis. Plant Signaling & Behavior 5 (11): 1–4.
 
Li R, Xiong G, Zhang B, and Zhou Y. (2010) Rice plants response to the disruption of OsCSLD4 gene. Plant Signaling & Behavior 5 (2): 1–4.
 
Tang C, Huang D, Yang J, Liu S, Sakr S, Li H, Zhou Y, and Qin Y. (2010) The sucrose transporter HbSUT3 plays an active role in sucrose loading to laticifer and rubber productivity in exploited trees of Hevea brasiliensis (rara rubber tree). Plant, Cell Environ. 33:1708–1720.
 
2009:
 
Li M, Xiong G, Li R, Cui J, Tang D, Zhang B, Pauly M, Cheng Z, and Zhou Y. (2009) Rice cellulose synthase-like D4 is essential for normal cell wall biosynthesis and plant growth. Plant J. 60: 1055–1069.
 
Zhang B, Deng L, Qian Q, Xiong G, Zeng D, Li R, Guo L, Li J, and Zhou Y. (2009) A missense mutation in the transmembrane domain of CESA4 affects protein abundance in the plasma membrane and results in abnormal cell wall biosynthesis in rice. Plant Mol. Biol. 71: 509–524. (Cover story)
 
Zhou Y, Li S, Qian Q, Zeng D, Zhang M, Guo L, Liu X, Deng L, Liu X, and Li J. (2009) BC10, a DUF266-containing and golgi-located type II membrane protein, is required for cell wall biosynthesis in rice (Oryza sativa L.). Plant J. 57, 446–462.
 
Before 2008:
 
Qi J, Qian Q, Bu Q, Li S, Chen Q, Sun J, Liang W, Zhou Y, Chu C, Li X, Ren F, Palme K, Zhao B, Chen J, Chen M, and Li C (2008). Mutation of the rice NARROW LEAF1 gene, which encodes a novel protein, affects vein patterning and polar auxin transport. Plant Physiology, 147, 1947–1959.
 
Lu, S., Zhou, Y., Li, L., and Chiang V. (2006) Distinct Roles of Cinnamate 4-hydroxylase Genes in Populus. Plant Cell Physiology, 47, 905–914.
 
Dai Y, Wang H Z, Li B H, Huang J, Liu X F, Zhou Y, Mou Z H, and Li J Y. (2005) Increased expression of MAP KINASE KINASE7 causes deficiency in polar auxin transport and leads to plant architectural abnormality in Arabidopsis. Plant Cell, 18, 308–320.
 
Zhao H Y, Liu J, Shiyou L, Zhou Y, Wei J H, Song Y R, and T Wang. (2005) Isolation and functional characterization of a cinnamate 4-hydroxylase promoter from populus tomentosa. Plant Science, 168, 1157–1162.
 
Li Y 1, Qian Q1, Zhou Y1, Yan M, Sun L, Zhang M, Fu Z, Wang Y, Han B, Pang X, Chen M, Li J. (2003) BRITTLE CULM1, which encodes a COBRA-like protein, affects the mechanical properties of rice plants. Plant Cell, 15, 2020–2031. (1Contribute equally)
 
Li L G, Zhou Y, Cheng X F, Sun J Y, Marita J M, Ralph J, Chiang V. (2003) Combinatorial modification of multiple lignin traits in trees through multigene cotransformation. Proc. Natl. Acad. Sci. USA. 100, 4939–4944.