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Plant and Cell Physiology Advance Access originally published online on May 15, 2007
Plant and Cell Physiology 2007 48(6):843-855; doi:10.1093/pcp/pcm055
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© The Author 2005. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oxfordjournals.org

Xyloglucan Endo-transglycosylase (XET) Functions in Gelatinous Layers of Tension Wood Fibers in Poplar—A Glimpse into the Mechanism of the Balancing Act of Trees

Nobuyuki Nishikubo1,5,8, Tatsuya Awano1,6,8, Alicja Banasiak1,7, Veronica Bourquin1, Farid Ibatullin2, Ryo Funada3, Harry Brumer2, Tuula T. Teeri2, Takahisa Hayashi4, Björn Sundberg1 and Ewa J. Mellerowicz1,*

1Department of Forest Genetics and Plant Physiology, Umea Plant Science Center, SLU, S901 83 Umea, Sweden
2Department of Biotechnology, Royal Insitute of Technology (KTH), S-106 91 Stockholm, Sweden
3Faculty of Agriculture, Tokyo University of Agriculture and Technology, Fuchu-Tokyo 183-8509 Japan
4Research Institute for Sustainable Humanosphere, Kyoto University, Kyoto, 611-0011 Japan

*Corresponding author: E-mail, ewa.mellerowicz{at}genfys.slu.se; Fax, +46-90-786-8165.


  Abstract

Tension wood is a specialized tissue of deciduous trees that functions in bending woody stems to optimize their position in space. Tension wood fibers that develop on one side of the stem have an increased potency to shrink compared with fibers on the opposite side, thus creating a bending moment. It is believed that the gelatinous (G) cell wall layer containing almost pure cellulose of tension wood fibers is pivotal to their shrinking. By analyzing saccharide composition and linkage in isolated G-layers of poplar, we found that they contain some matrix components in addition to cellulose, of which xyloglucan is the most abundant. Xyloglucan, xyloglucan endo-transglycosylase (XET) activity and xyloglucan endo-transglycosylase/hydrolase (XTH) gene products were detected in developing G-layers by labeling using CCRC-M1 monoclonal antibody, in situ incorporation of XXXG-SR and the polyclonal antibody to poplar PttXET16-34, respectively, indicating that xyloglucan is incorporated into the G-layer during its development. Moreover, several XTH transcripts were altered and were generally up-regulated in developing tension wood compared with normal wood. In mature G-fibers, XTH gene products were detected in the G-layers while the XET activity was evident in the adjacent S2 wall layer. We propose that XET activity is essential for G-fiber shrinking by repairing xyloglucan cross-links between G- and S2-layers and thus maintaining their contact. Surprisingly, XTH gene products and XET activity persisted in mature G-fibers for several years, suggesting that the enzyme functions after cell death repairing the cross-links as they are being broken during the shrinking process.

Keywords: Gravity responses - Populus - Reaction wood - Secondary wall - Wood formation - Xylogenesis

Abbreviations: ADF6, actin depolymerizing factor 6; CW, compression wood; DMSO, dimethylsulfoxide; elF4A-8, eukaryotic initiation factor 4A-8; EST, expressed sequence tag; FITC, fluorescein isothiocyanate; G, gelatinous; NW, normal wood; OW, opposite wood; TW, tension wood; UBQ4, polyubiquitin 4; XET, xyloglucan endo-transglycosylase; XG, xyloglucan; XTH16, xyloglucan endo-transglycosylase/hydrolase of family GH16; XXXG-SR, sulforhodamine-labeled xylogluco-oligosaccharide

5Present address: RIKEN Plant Science Center, 1-7-22 Suehiro-cho Tsurumi-ku, Yokohama, Kanagawa, 230-0045 Japan.

6Present address: Laboratory of Tree Cell Biology, Division of Forest and Biomaterials Science, Graduate School of Agriculture, Kyoto University, Kyoto, 606-8502 Japan.

7Present address: Institute of Plant Biology, University of Wroclaw, Kanonia 6/8, 50-328, Wroclaw, Poland.

8These authors contributed equally to this work.

(Received March 1, 2007; Accepted May 8, 2007)
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