Abstract
Here we discuss the formation of phyllotactic patterns in the shoot apical meristem (SAM) of plants, where the spatial distribution of the phytohormone auxin determines phyllotaxis in a domain that is growing and changing in time. We assume that the concentration of auxin modifies the mechanical properties of the domain and that the mechanical stress field in the SAM orients the flux of auxin. To study this problem we propose a mechanism for pattern formation in growing domains with variable curvature. The dynamics of chemicals is modeled by a reaction-diffusion system that produces a three dimensional pattern of chemical concentrations that changes the stress field in the domain while growing. The growth process is modeled by a phase-field order parameter which determines the location of the boundaries of the domain. This field is coupled to the chemical concentration through a curvature term that affects the local mechanical stress in the domain. The local stress changes in turn modify the chemical patterns. Our model constitutes a useful and novel approach in theoretical biology, as many developmental processes in organisms seem to be affected by the changes of curvature, size, mechanical stress and other physical aspects. Several patterns seen in many plants are reproduced under certain conditions by our model.
Highlights
Phyllotaxis is the arrangement of repeated plant organs such as leaves, floral structures, ribs in cacti or scales in a pine
The first whorl emerges at the basis of the dome, which is the region where the curvature of the domain changes more abruptly (Fig 2C). This is in agreement with experimental observations which indicate that the primordia emerge exactly where the curvature of the shoot apical meristem (SAM) dome changes [78]
The domain evolves according to the minimization of a free energy that is constructed on the basis of the bending energy of a surface, and as a result of the influence that a chemical substance has on its mechanical properties
Summary
Phyllotaxis is the arrangement of repeated plant organs such as leaves, floral structures, ribs in cacti or scales in a pine. This geometrical patterning has fascinated minds for centuries. The scientific importance of studying the form of living organisms was pertinently pointed out by Darcy Thompson [1]. Phyllotactic patterns can be obtained with very simple space filling physical rules, as in foams [2] and Benard convection cells [3].
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