Auxin

Auxins (plural of auxin /ˈɔːksɪn/) are a class of plant hormones (or plant-growth regulators) with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essential for plant body development. The Dutch biologist Frits Warmolt Went first described auxins and their role in plant growth in the 1920s. Kenneth V. Thimann became the first to isolate one of these phytohormones and to determine its chemical structure as indole-3-acetic acid (IAA). Went and Thimann co-authored a book on plant hormones, Phytohormones, in 1937.

Auxins were the first of the major plant hormones to be discovered. They derive their name from the Greek word αὔξειν (auxein – 'to grow/increase'). Auxin is present in all parts of a plant, although in very different concentrations. The concentration in each position is crucial developmental information, so it is subject to tight regulation through both metabolism and transport. The result is the auxin creates "patterns" of auxin concentration maxima and minima in the plant body, which in turn guide further development of respective cells, and ultimately of the plant as a whole.

The (dynamic and environment responsive) pattern of auxin distribution within the plant is a key factor for plant growth, its reaction to its environment, and specifically for development of plant organs (such as leaves or flowers). It is achieved through very complex and well-coordinated active transport of auxin molecules from cell to cell throughout the plant body—by the so-called polar auxin transport. Thus, a plant can (as a whole) react to external conditions and adjust to them, without requiring a nervous system. Auxins typically act in concert with, or in opposition to, other plant hormones. For example, the ratio of auxin to cytokinin in certain plant tissues determines initiation of root versus shoot buds.

On the molecular level, all auxins are compounds with an aromatic ring and a carboxylic acid group. The most important member of the auxin family is indole-3-acetic acid (IAA), which generates the majority of auxin effects in intact plants, and is the most potent native auxin. And as native auxin, its equilibrium is controlled in many ways in plants, from synthesis, through possible conjugation to degradation of its molecules, always according to the requirements of the situation. Auxin can act in a heat sensitive manner in many situations, which will in turn effect a plant's phenotypic development.

Some synthetic auxins, such as 2,4-D and 2,4,5-trichlorophenoxyacetic acid (2,4,5-T), are sold as herbicides. Broad-leaf plants (dicots), such as dandelions, are much more susceptible to auxins than narrow-leaf plants (monocots) such as grasses and cereal crops, making these synthetic auxins valuable as herbicides.

In 1881, Charles Darwin and his son Francis performed experiments on coleoptiles, the sheaths enclosing young leaves in germinating grass seedlings. The experiment exposed the coleoptile to light from a unidirectional source, and observed that they bend towards the light. By covering various parts of the coleoptiles with a light-impermeable opaque cap, the Darwins discovered that light is detected by the coleoptile tip, but that bending occurs in the hypocotyl. However the seedlings showed no signs of development towards light if the tip was covered with an opaque cap, or if the tip was removed. The Darwins concluded that the tip of the coleoptile was responsible for sensing light, and proposed that a messenger is transmitted in a downward direction from the tip of the coleoptile, causing it to bend.

In 1910, Danish scientist Peter Boysen Jensen demonstrated that the phototropic stimulus in the oat coleoptile could propagate through an incision. These experiments were extended and published in greater detail in 1911 and 1913. He found that the tip could be cut off and put back on, and that a subsequent one-sided illumination was still able to produce a positive phototropic curvature in the basal part of the coleoptile. He demonstrated that the transmission could take place through a thin layer of gelatin separating the unilaterally illuminated tip from the shaded stump. By inserting a piece of mica he could block transmission in the illuminated and non-illuminated side of the tip, respectively, which allowed him to show that the transmission took place in the shaded part of the tip. Thus, the longitudinal half of the coleoptile that exhibits the greater rate of elongation during the phototropic curvature, was the tissue to receive the growth stimulus.

In 1911, Boysen Jensen concluded from his experimental results that the transmission of the phototropic stimulus was not a physical effect (for example due to a change in pressure) but serait dû à une migration de substance ou d'ions (was caused by the transport of a substance or of ions). These results were fundamental for further work on the auxin theory of tropisms.