Rolf Rutishauser, University of Zurich, Zurich, Switzerland
Peter Peisl, University of Zurich, Zurich, Switzerland
Published online: April 2001
DOI: 10.1038/npg.els.0002057
Phyllotaxy (phyllotaxis) is the mode of arrangement of leaves, scales, or bracts with flowers along the plant stem. Phyllotaxy research deals with the study of biological pattern formation (morphogenesis); it answers questions such as what the shoot apical meristem (SAM) does and how it does it.
Keywords: Fibonacci systems; Lucas systems; pattern formation; phyllotaxy theories; shoot apical meristem; whorls
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Figure 1.
Shoot tip of a member of the carnation family (Honkenya peploides), showing decussate phyllotaxy, i.e. crossed leaf pairs. Leaf primordia arise pairwise and opposite each other on the flank of the shoot apical meristem (diameter 150 μm). |
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Figure 2.
Shoot tip of a shrubby plantain (Plantago webbii) with tetramerous whorls. Four leaf primordia are equally distributed around the shoot apical meristem (diameter 80 μm). The leaves of the next older tetramerous whorl are removed except one. |
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Figure 3.
Shoot tip of a yellow stonecrop (Sedum sexangulare), showing spiral pattern according to Lucas phyllotaxy with divergence angle approaching 99.5°. Four successively formed leaf primordia surround the shoot apical meristem (diameter 80 μm). |
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Figure 4.
Rosette (diameter 30 cm) of Canarian house‐leek (Aeonium tabuliforme) representing a Fibonacci system with 8 left‐turning contact parastichies (two of them marked with red stickers) and 13 right‐turning contact parastichies (three of them marked with orange stickers). Six leaves (1–6) close to the centre are marked with yellow stickers according to the genetic spiral (divergence angle approaching 137.5°). |
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Figure 5.
Central portion of sunflower (Helianthus annuus) capitulum, with 29 left‐turning contact parastichies and 47 right‐turning contact parastichies. One of each set is marked with black dots. The contact parastichies [29 + 47] are members of the Lucas sequence 1, 3, 4, 7, 11, …. Lucas phyllotaxy occurs in ∼5% of all sunflower capitula, whereas Fibonacci systems are much more frequent. |
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Figure 6.
Pine‐cone (Pinus nigra) seen from the base, representing bijugate phyllotaxy, another rare spiral system. There are paired scales that are opposite each other or nearly so (labelled with the same even numeral). Consecutive leaf pairs (0–2–4–6– …) are twisted ∼69°, i.e. the half Fibonacci angle. This rare spiral pattern shows contact parastichies [6 + 10], which are members of the sequence 4, 6, 10, 16, …. |
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Figure 7.
All spiral phyllotaxes show chirality. For example, there are as many right‐handed as left‐handed pine‐cones on a tree (Pinus sp.). The 8 contact parastichies can run from bottom right to top left or from bottom left to top right. |
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Figure 8.
Surface of an aberrant pineapple (Ananas comosus), peeled off and unrolled. There are eight parastichies running from bottom left to top right. Two of the 12 parastichies running from bottom right to top left fuse into one (green dot); thus there are only 11 parastichies higher up. Normal pineapples show contact parastichy patterns of [8 + 13]. |
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Figure 9.
Computer‐simulated Fibonacci system illustrating cross‐section of shoot tip with tightly packed leaves. One sample of each recognizable parastichy set is marked with a different colour: 13 (green) and 21 (blue) are the most conspicuous contact parastichy sets. Less obvious are parastichy sets along which the leaves have (nearly) no contact: 8 (yellow) and 34 (pink). |
Further Reading
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Carpenter R, Copsey L, Vincent C et al. (1995) Control of flower development and phyllotaxy by meristem identity genes in Antirhinum. The Plant Cell 7(12): 2001–2011.
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