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

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).

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.

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).

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°).

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.

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, ….

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.

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].

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

Callos JD and Medford JI (1994) Organ positions and pattern formation in the shoot apex. Plant Journal 6: 1–7.

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.

Cottignies A (1994) Phyllotaxie. Acta Botanica Gallica 141: 5–14.

Endress PK (1994) Diversity and Evolutionary Biology of Tropical Flowers. Cambridge: Cambridge University Press.

Fleming AJ, Caderas D, Wehrli E, McQueen‐Mason S and Kuhlemeier C (1999) Analysis of expansin‐induced morphogenesis on the apical meristem of tomato. Planta 208: 166–174.

Gola E (1996) Phyllotaxis diversity in Lycopodium clavatum and Lycopodium annotinum. Acta Societatis Botanicorum Poloniae 65: 235–247.

Green PB (1999) Expression of pattern in plants: combining molecular and calculus‐based biophysical paradigms. American Journal of Botany 86: 1059–1076.

Hofmeister W (1868) Allgemeine Morphologie der Gewächse. Handbuch der physiologischen Botanik, vol. 1, pp. 405–664. Leipzig: Engelmann.

Jean RV (1994) Phyllotaxis. A Systemic Study in Plant Morphogenesis. Cambridge: Cambridge University Press.

Jean RV and Barabé D (eds) (1998) Symmetry in Plants. Singapore: World Scientific.

Kirchoff BK and Rutishauser R (1990) The phyllotaxy of Costus (Costaceae). Botanical Gazette 151: 88–105.

Kwiatkowska D (1999) Formation of pseudowhorls in Peperomia verticillata (L.) A. Dietr. shoots exhibiting various phyllotactic patterns. Annals of Botany 83: 675–685.

Lyndon RF (1998) The Shoot Apical Meristem: Its Growth and Development. New York: Cambridge University Press.

McConnell JR and Barton MK (1998) Leaf polarity and meristem formation in Arabidopsis. Development 125: 2935–2942.

Pogany JA, Simon EJ, Katzman RB et al. (1998) Identifying novel regulators of shoot meristem development. Journal of Plant Research 111: 307–313.

Prusinkiewicz P and Lindenmayer A (1990) The Algorithmic Beauty of Plants. New York: Springer‐Verlag.

Rutishauser R (1999) Polymerous leaf whorls in vascular plants: developmental morphology and fuzziness of organ identity. International Journal of Plant Sciences 160 (6, supplement): S81–S103.

Scanlon MJ (1998) Force fields and phyllotaxy: an old model comes of age. Trends in Plant Science 3: 413–414.

Theissen G and Saedler H (1998) Molecular architects of plant body plans. Progress in Botany 59: 227–256.

Tsukaya H (1998) Genetic evidence for polarities that regulate leaf morphogenesis. Journal of Plant Research 111: 113–119.

van der Linden FMJ (1996) Phyllotactic Patterns for Domes. Patterns and Morphology of Building Elements for Close‐packing of Dome Surfaces after the Example of Phyllotaxis and Morphogenesis of Primordia on the Apex of Composites. De Lier: Academisch Boeken Centrum.

Veit B, Briggs SP, Schmidt RJ, Yanofsky MF and Hake S (1998) Regulation of leaf initiation by the terminal ear 1 gene of maize. Nature 393: 166–168.

Zagórska‐Marek B (1994) Phyllotaxic diversity in Magnolia flowers. Acta Societatis Botanicorum Poloniae 63: 117–137.

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How to Cite close
Rutishauser, Rolf, and Peisl, Peter(Apr 2001) Phyllotaxy. In: eLS. John Wiley & Sons Ltd, Chichester. [doi: 10.1038/npg.els.0002057]