Phyllotaxis
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In botany, phyllotaxis or phyllotaxy is the arrangement of the leaves on the stem of a plant.
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[edit] Pattern structure
The basic patterns are alternate, opposite, whorled or spiral. With an alternate pattern, leaves switch from side to side. An alternate distichous phyllotaxis means that each leaf growing at a single node is disposed in a single rank along the branch (such as in grasses). In an opposite pattern, two leaves grow in opposite directions from the same node. In an opposite pattern, if successive leaf pairs are perpendicular, this is called decussate. A whorled pattern consists of three or more leaves at each node. An opposite leaf pair can be thought of as a whorl of two leaves. A whorl can occur as a basal structure where all the leaves are attached at the base of the shoot and the internodes are small or nonexistent. A basal whorl with a large number of leaves spread out in a circle is called a rosette. A multijugate pattern is a spiral composed of whorls. The pattern has also been observed to emerge in at least one animal cell (the red blood cell), during processes that perturb cellular fluid dynamics [1].
[edit] Repeating spiral
A repeating spiral can be represented by a fraction describing the angle of windings leaf per leaf.
Alternate leaves will have an angle of 1/2 of a full rotation. In beech and hazel the angle is 1/3, in oak and apricot it is 2/5, in poplar and pear it is 3/8, and in willow and almond the angle is 5/13.[2] The numerator and denominator normally consist of a Fibonacci number and its second successor. The number of leaves is sometimes called rank, in the case of simple Fibonacci ratios, because the leaves line up in vertical rows. With larger Fibonacci pairs, the pattern becomes complex and non-repeating. This tends to occur with a basal configuration. Examples can be found in composite flowers and seed heads. The most famous example is the sunflower head. This phyllotactic pattern creates an optical illusion of criss-crossing spirals. In the botanical literature, these designs are described by the number of counter-clockwise spirals and the number of clockwise spirals. These also turn out to be Fibonacci numbers. In some cases, the numbers appear to be multiples of Fibonacci numbers because the spirals consist of whorls.
Leonardo da Vinci was the first to suggest that the adaptive advantage of the Fibonacci pattern is to maximize exposure to dew. Current thinking supports this interpretation. Phyllotactic architecture optimizes access to moisture, rainfall and sunlight.
[edit] Determination
The pattern of leaves on a plant is controlled by the plant hormone auxin.[3]
[edit] History
Insight into the mechanism had to wait until Wilhelm Hofmeister proprosed a model in 1868. The process begins with two primordia, nascent leaves, forming on opposite sides of the shoot. A third new leaf then erupts between them. Each old leaf pushes the upstart away. The golden angle is the blind result of this jostling. Since three golden arcs add up to slightly more than enough to wrap a circle, one of the old leaves overlaps and is pushed out in a radial line, like a rocket that escapes earth orbit. This vacates space in the inner generative spiral for a new leaf to form. The generative spiral should not be confused with the clockwise and counter-clockwise spirals that emerge in densely packed plant structures. Spirals are discerned by tracing the path from leaf to neighboring leaf and each leaf acquires new neighbors as it shoots out from the center and its sister primordia. New leaves move into the spaces opened as old leaves diverge. Phi is an irrational number and this guarantees that no two leaves ever follow the same radial line from center to edge.
In modern times, researchers such as Snow and Snow[citation needed] have continued these lines of inquiry. Computer modeling and morphological studies have confirmed and refined Hoffmeister's ideas. Questions remain about the details. Botanists are divided on whether the control of leaf migration depends on chemical gradients among the primordia or purely mechanical forces. Lucas rather than Fibonacci numbers have been observed in a few plants and occasionally the leaf positioning appears to be random.
[edit] See also
[edit] Notes
- ^ J.T.Lofthous Phyllotaxis Patterns in Animal Cells: A Dynamic Mechanism for Red Cell Echinocytosis and Programmed Cell Death.[1]
- ^ Coxeter, H. S. M. (1961), Introduction to geometry, Wiley, pp. 169
- ^ http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=1692983 Philos Trans R Soc Lond B Biol Sci. 2002 June 29; 357(1422): 737–747. The shoot apical meristem: the dynamics of a stable structure. Jan Traas and Teva Vernoux
[edit] References
- F.M.J. van der Linden
- Frank M.J. van der Linden: Creating Phyllotaxis, The Stack-and-Drag model, in Mathematical Biosciences, NY 1996
- Frank M.J. van der Linden: Creating Phyllotaxis from Seed to Flower, in Symmetry in Plants, Jean & Barabé eds., World Scientific, Singapore 1998
[edit] External links
- Fibonacci Numbers and the Golden Section
- Spiral Phyllotaxis Pattern in an Animal Cell: A Fluid- Driven Mechanism for Red Cell Echinocytosis and Programmed Cell Death- a paper by Dr. Juanita Lofthouse presenting the first example of an animal cell exhibiting Phyllotaxis patterns in response to changes in cell metabolic rate and other dynamic perturbations
- Phyllotaxis as a Dynamical Self Organizing Process
- Eric W. Weisstein, Phyllotaxis at MathWorld.
- Phyllotaxis Spirals and Phyllotaxis Spirals in 3D by Stephen Wolfram, The Wolfram Demonstrations Project.
- Phyllotaxis: An Interactive Site for the Study of Plant Pattern Formation
- PhaseLab
