Vascular Cambium—Between the Hammer and the Anvil: A Tensile Stress Hypothesis on the Mechanism of Radial Growth of Broadleaved Trees

Abstract

The vascular cambium is the main lateral meristem responsible for the secondary growth of trees. There are a number of explicit and implicit assumptions behind this statement which allow questions to be raised about the mechanism underlying the radial growth of trees. Based on the hypothesis of the diurnal strains of plant organs, it is anticipated that the process of radial growth can be understood as an adaptation to the cyclically changing mechanical stress in the radial direction generated by the phloem during the 24 h day cycle. This qualitative hypothesis treats cambium as a tissue subjected to nighttime stretching and daytime compression in the radial direction. The proposed osmo-mechanical hypothesis of the radial growth of vascular cambium links the daily change in water status and the considerable daily strains in the xylem and phloem with the radial net expansion of a tree trunk. We highlight transpiration as a major factor influencing the secondary growth of woody plants. Thus, we indirectly relate all the biotic (e.g., insect infestation, fungi infections, injuries, shadowing, intra- and interspecies competition, parasitism, symbiosis, etc.) and abiotic (e.g., humidity, water availability, wind, injuries, shadowing, day length in a vegetative season, altitude, temperature, insolation, etc.) processes influencing transpiration with radial growth. In the proposed hypothesis, we also infer that differences in the strains in phloem and xylem are the direct source of tensile stress, tensile stress relaxation, compressive stress, and compressive stress relaxation in the vascular cambium. This sequence appears to be crucial in understanding both the process of the radial growth of trees and the formation of differential wood patterns, within the same genotype as well as in different genotypes. It also provides arguments for the discussion on the mechanisms regulating processes in the vascular cambium. It points out the important role of the variable mechanical stresses in the radial, circumferential, and axial directions and their interference in the development of this lateral meristem. Thus, this hypothesis supports the concept of the epigenetic and systemic regulation of intrinsic wood patterns and tree forms by environmental factors. The hypothesis is focused exclusively on broadleaved trees and symplastic growth. This limitation of the scope is due to a concern for clarity. In this form, the hypothesis provides an alternative explanation for a pure process of radial growth and paves the way for a better interpretation of such phenomena as earlywood and latewood formation. At the same time, this approach to the vascular cambium provides answers to many questions related to the generation of the mechanical conditions necessary for the occurrence of intrusive growth between tangential cell walls; this is of fundamental importance for fusiform initials readjustment, vessel element and fibre formation, ring-porous wood formation, etc.