Gravistimulation of tree stems affects wood development by unilaterally inducing wood with modified properties, called reaction wood. Commonly, it also stimulates cambial growth on the reaction wood side. Numerous experiments involving applications of indole-3-acetic acid (IAA) or IAA-transport inhibitors have suggested that reaction wood is induced by a redistribution of IAA around the stem. However, in planta proof for this model is lacking. Therefore, we have mapped endogenous IAA distribution across the cambial region tissues in both aspen (Populus tremula, denoted poplar) and Scots pine (Pinus sylvestris) trees forming reaction wood, using tangential cryosectioning combined with sensitive gas chromatography-mass spectrometry analysis. Moreover, we have documented the kinetics of IAA during reaction wood induction in these species. Our analysis of endogenous IAA demonstrates that reaction wood is formed without any obvious alterations in IAA balance. This is in contrast to gravitropic responses in roots and shoots where a redistribution of IAA has been documented. It is also of interest that cambial growth on the tension wood side was stimulated without an increase in IAA. Taken together, our results suggest a role for signals other than IAA in the reaction wood response, or that the gravitational stimulus interacts with the IAA signal transduction pathway.
This thesis considers aspects of the regulation of growth rate and fibre properties in forest trees. These properties are both genetically determined and influenced by environmental stimuli. Induction of reaction wood is an environmentally induced process involving changes in growth rate and fibre properties that can be readily studied. Plant hormones are signalling agents that play important roles in the initiation and coordination of wood formation; in this thesis the plant hormones auxin and ethylene were investigated using gas chromatography/mass spectrometry (GC/MS). A novel MS technique for measuring the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC) in minute amounts of plant tissue was developed. Ethylene is often connected to stress responses in plants, and ethylene evolution is increased when reaction wood is formed. Here it is demonstrated that this increase is regulated by ACC oxidase, the enzyme catalysing the last step in the ethylene biosynthetic pathway. This is in contrast to most of the earlier findings that tended to indicate that ethylene production directly reflects the availability of ACC. Although ethylene is strongly up-regulated during reaction wood formation, its role in modulating the growth rate and fibre properties remains unknown. Further, it is demonstrated that reaction wood in both poplar (Populus tremula L.) and pine (Pinus sylvestris L.) is formed without changes in auxin concentration in the cambial tissues. This suggests that the previously held assumption that the difference in auxin concentration is key factor in the induction of reaction wood is unsound. Further, auxin concentrations were compared in hybrid aspen trees (Populus tremula L. x tremuloides Michx.) growing vertically at different growth rates. These trees showed good correlations between auxin levels and growth rates. The growth rate was mediated by increases in the cell cycling rate rather than in the width of the cell division zone. Thus, the growth rate in poplar was correlated to auxin levels in normal wood formation, but not during reaction wood formation.
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