Vascular phyllotaxis transition and an evolutionary mechanism of phyllotaxis

Leaves of vascular plants are arranged regularly around stems, a phenomenon known as phyllotaxis. A constant angle between two successive leaves is called divergence angle. On the one side, the divergence angle $\alpha_0$ of an initial pattern of leaf primordia at a shoot apex is most commonly an irrational number of about 137.5 degrees, called limit divergence. On the other side, the divergence $\alpha$ of a final pattern of leaf traces in the vascular system of a mature stem is expressed in terms of a sequence of rational numbers, 1/2, 1/3, 2/5, 3/8, 5/13, 8/21, called phyllotactic fractions. The mathematical relationship between the initial divergence $\alpha_0$, the final divergence $\alpha$, and the number of internodes traversed by the leaf traces $n_c$ is investigated by means of a theoretical model of vascular phyllotaxis. It is shown that continuous changes of the trace length $n_c$ induce transitions between the fractional orders in the vascular structure. The vascular phyllotaxis transition suggests an evolutionary mechanism for the phenomenon of phyllotaxis. To provide supporting evidence for the model and mechanism, available experimental results for fossil remains of Lepidodendron and the vascular structure of Linum and Populus are analyzed with the model.