Growth stress controls negative gravitropism in woody plant stems

Abstract. In the shoots of woody plant species, reaction-wood fibers are formed on the upper or lower side of the secondary xylem of a leaning trunk or branch wherever large, internal growth stress is generated. Negative gravitropic movement in woody plant stems is proposed to be the result of growth stress generated in the reaction-wood tissue. This study examines the interaction between bending moment due to increasing self-weight and recovery moment resulting from asymmetric growth stress, and tests a hypothesis that describes the relationship based on the structural mechanics "beam theory". Simulations of observed tree branch morphology of Magnolia kobus DC., Juniperus chinensis L., Abies saccharinensis Fr. Schum., and Prunus spachiana Kitamura f. spachiana cv. Plenarosea showed that (i) the growth stress generated in the reaction wood is sufficient to counteract the gravitropic response to increasing self-weight, and (ii) the specific directional angle of the shoot apex or preferred angle of the elongation zone plays an important role in controlling the spatial shape of the branch stem that is peculiar to plant species with large growth stress generated in the reaction-wood tissue.

[1]  Henri Baillères,et al.  Tree biomechanics : growth, cumulative prestresses, and reorientations , 1994 .

[2]  T. Fourcaud,et al.  Mechanical analysis of the form and internal stresses of a growing tree by the finite element method , 1995 .

[3]  T. E. Timell Compression Wood in Gymnosperms , 1986 .

[4]  H. Yamamoto,et al.  Generation mechanism of growth stresses in wood cell walls: roles of lignin deposition and cellulose microfibril during cell wall maturation , 1998, Wood Science and Technology.

[5]  B. Gartner,et al.  Lean in red alder (Alnus rubra) : growth stress, tension wood, and righting response , 1996 .

[6]  J. D. Boyd,et al.  Tree growth stresses — Part V: Evidence of an origin in differentiation and lignification , 1972, Wood Science and Technology.

[7]  Tissue stresses and their graviresponsive changes in stems of Reynoutria japonica Houtt , 1996 .

[8]  Hiroyuki Yamamoto,et al.  Growth stresses in tension wood: role of microfibrils and lignification , 1994 .

[9]  Dr. Robert R. Archer,et al.  Growth Stresses and Strains in Trees , 1987, Springer Series in Wood Science.

[10]  P Castéra,et al.  Growth patterns and bending mechanics of branches , 1991, Trees.

[11]  J. Palmer Epinasty, Hyponasty, and Related Topics , 1985 .

[12]  R. Hangarter,et al.  Gravity, light and plant form. , 1997, Plant, cell & environment.

[13]  Masato Yoshida,et al.  Generation process of growth stresses in cell walls II. Growth stresses in tension wood. , 1990 .

[14]  W. Silk DESCRIPTIONS OF DEVELOPMENT , 1984 .

[15]  Masato Yoshida,et al.  Relationship between growth rate and growth stresses in Paraserianthes falcataria grown in Indonesia. , 2000 .

[16]  Masato Yoshida,et al.  Growth stresses and strains in Acacia mangium , 1999 .

[17]  R. R. Archer,et al.  On the distribution of tree growth stresses — Part I: An anisotropic plane strain theory , 2004, Wood Science and Technology.

[18]  Masato Yoshida,et al.  Tension wood and growth stress induced by artificial inclination in Liriodendron tulipifera Linn. and Prunus spachiana Kitamura f. ascendens Kitamura , 2000 .

[19]  R. Archer,et al.  Apical control of branch movements in white pine: Compression wood action , 1982, Wood Science and Technology.

[20]  S. Wyatt,et al.  Growth dynamics and cytoskeleton organization during stem maturation and gravity-induced stem bending in Zea mays L. , 1998, Planta.

[21]  Mechanics of the Compression Wood Response: II. On the Location, Action, and Distribution of Compression Wood Formation. , 1973, Plant physiology.

[22]  A. Sievers,et al.  Tissue stresses in organs of herbaceous plants. II: Determination in three dimensions in the hypocotyl of sunflower , 1995 .

[23]  M. Fournier,et al.  Growth stress patterns in tree stems , 1990, Wood Science and Technology.

[24]  K. Tanaka,et al.  RESIDUAL STRESSES—III: Science and Technology , 1992 .

[25]  Zygmunt Hejnowicz,et al.  Graviresponses in herbs and trees: a major role for the redistribution of tissue and growth stresses , 1997, Planta.

[26]  Y. Masuda,et al.  Geotropic response of cucumber hypocotyls , 1974 .

[27]  R. Archer,et al.  Mechanics of the Compression Wood Response , 2022 .

[28]  D. Cosgrove Cellular mechanisms underlying growth asymmetry during stem gravitropism , 1997, Planta.