title: Seasonal and clonal variation in cellulose microfibril orientation during cell wall formation of tracheids in Cryptomeria japonica.
abstract: To investigate the biological mechanism by which trees control the changes in microfibril (MF) orientation among secondary cell wall layers of conifer tracheids, we studied seasonal variation in the orientation of newly deposited MFs during tracheid cell wall development in Japanese cedar (Cryptomeria japonica D. Don) trees growing in Central Japan (36°36'N, 140°39'E). Sample blocks were repeatedly collected from four 16-year-old clones of different origins during the growing season of 2010 to investigate the hypotheses that changes in cellulose MF orientation between wall layers exhibited seasonal and clonal differences. The progressive change in the orientation of newly deposited MFs on the primary and secondary cell wall layers of tracheids was detected by field-emission-scanning electron microscopy. Tracheid production and differentiation was studied by light microscopy. We observed a decreasing trend in the orientation of deposited MFs from earlywood to latewood in the S2 and S1 layers, where MFs appeared in a Z-helix. In contrast, no seasonal pattern in the orientation of the MFs in the S-helix was observed. Minor clonal variation was observed in the phenology of tracheid production and differentiation. We concluded that a seasonal decreasing trend in the orientation of the MFs in the Z-helix in S1 and S2 was present, whereas the MFs in other layers exhibited minor random variations. Thus, the orientation of the MFs in S2 was affected by seasonal factors, whereas the MFs in other layers were more intrinsically controlled. The within-ring variations in the MF orientation and thus the resulting average MF angle might also be related to genotypic differences in the tracheid production and differentiation rate. However, our results do not exclude other intrinsic and environmental regulations in the change in MF orientation, which remains a topic for future studies.
authors.affiliation: Finnish Forest Research Institute, Vantaa Research Unit, PO Box 18, FI-01301 Vantaa, Finland tuula.jyske@metla.fi.
authors.forename: Tuula
authors.initials: T
authors.lastname: Jyske
authors.affiliation: Forestry and Forest Products Research Institute, 1 Matsunosato, Tsukuba, Ibaraki 305-8687, Japan.
authors.forename: Takeshi
authors.initials: T
authors.lastname: Fujiwara
authors.affiliation: Forestry and Forest Products Research Institute, 1 Matsunosato, Tsukuba, Ibaraki 305-8687, Japan.
authors.forename: Katsushi
authors.initials: K
authors.lastname: Kuroda
authors.affiliation: Forest Tree Breeding Center, Forestry and Forest Products Research Institute, 3809-1 Ishi, Hitachi, Ibaraki 319-1301, Japan.
authors.forename: Taiichi
authors.initials: T
authors.lastname: Iki
authors.affiliation: Forestry and Forest Products Research Institute, 1 Matsunosato, Tsukuba, Ibaraki 305-8687, Japan.
authors.forename: Chunhua
authors.initials: C
authors.lastname: Zhang
authors.affiliation: Finnish Forest Research Institute, Vantaa Research Unit, PO Box 18, FI-01301 Vantaa, Finland Forestry and Forest Products Research Institute, 1 Matsunosato, Tsukuba, Ibaraki 305-8687, Japan Forest Tree Breeding Center, Forestry and Forest Products Research Institute, 3809-1 Ishi, Hitachi, Ibaraki 319-1301, Japan.
authors.forename: Tuomas K
authors.initials: TK
authors.lastname: Jyske
authors.affiliation: Forestry and Forest Products Research Institute, 1 Matsunosato, Tsukuba, Ibaraki 305-8687, Japan.
authors.forename: Hisashi
authors.initials: H
authors.lastname: Abe
mesh_terms: D002473:Cell Wall; D002482:Cellulose / Q000378:metabolism*; D032485:Cryptomeria / Q000235:genetics / Q000502:physiology*; D005838:Genotype; D007564:Japan; D020894:Microfibrils / Q000378:metabolism*; D012621:Seasons; D014197:Trees / Q000502:physiology; D014934:Wood / Q000254:growth & development; D052584:Xylem / Q000166:cytology / Q000254:growth & development*
chemical_list: D002482:Cellulose
keywords: Japanese cedar; tracheid differentiation