Phyllotaxis studies published in German in the 1930s have reported intriguing regularity in the arrangement of incipient leaves on shoot apices of a wide variety of plant species. However, these studies have received little attention today, even though they provide a crucial evidence base for understanding this mathematical phenomena. Here I recapitulate the essential point by means of illustrative examples. It is emphasized that accurate control of apical divergence angle is at the heart of the numerical riddle of spiral phyllotaxis. The accurate patterning at the shoot apex has an unexpected evolutionary benefit of being optimally adaptive in the subsequent events of phyllotactic change to occur on an elongating shoot.
Divergence angles were measured for inflorescences of Helianthus annuus with several hundreds to more than a thousand disk flowers. Quantitative analysis showed that the angles are robustly fixed in the vicinity of the ideal golden angle 137.508° as accurately as ~0.001°. The mean deviation from the ideal value varies for each sample. The results have important implications for phyllotaxis models, which are discussed by referring to a necessary modification proposed by Roberts.
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