auxin

auxin, plant hormone that regulates the amount, type, and direction of plant growth. Auxins include both naturally occurring substances and related synthetic compounds that have similar effects. Auxins are found in all members of the plant kingdom. They are most abundantly produced in growth areas (meristem), e.g., root and shoot tips, but are also produced elsewhere, e.g., in the stems and leaves. The method of dispersal throughout the plant body is not yet fully understood. Auxins affect numerous plant processes, e.g., cell division and elongation, autumnal loss of leaves, and the formation of buds, roots, flowers, and fruit. They are also responsible for many forms of tropism. It is known that phototropism is due to the inhibition of auxins by light; the cells on that side of a plant exposed to light do not divide or grow as quickly as those on the shaded side, and thus the plant grows toward the light source. Auxins are widely used commercially to produce more vigorous growth, to promote flowering and fruiting and also root formation in plants not easily propagated by stem cuttings, to retard fruit drop, and to produce seedless varieties (e.g., of tomatoes) by parthenogenetic fruiting. Only minute amounts of auxins occur naturally, and synthetic auxins (e.g., 2,4-D) must be administered in carefully prescribed doses, since excessive concentration produces usually fatal abnormalities. However, different species of plants react to different amounts of auxins, a fact used to advantage as a method of weed control. The principal natural auxin is indoleacetic acid; other common but less frequent plant hormones include the gibberellins, lactones, and kinins.

Any of a group of hormones that regulate plant growth, particularly by stimulating cell elongation in stems and inhibiting it in roots. Auxins influence the growth of stems toward light (phototropism) and against the force of gravity (geotropism). Auxins also play a role in cell division and differentiation, fruit development, the formation of roots from cuttings, the inhibition of lateral branching, and leaf fall. The most important naturally occurring auxin is beta-indolylacetic acid.

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Auxins are a class of plant growth substance (often called phytohormone or plant hormone). Auxins play an essential role in coordination of many growth and behavioral processes in the plant life cycle.

Overview

Auxins derive their name from the Greek word auxano (to grow). They were the first of the major plant hormones to be discovered and are a major coordinating signal in plant development. Their pattern of active transport through the plant is complex. They typically act in concert with (or opposition to) other plant hormones. For example, the ratio of auxin to cytokinin in certain plant tissues determines initiation of root versus shoot buds. Thus a plant can (as a whole) react on external conditions and adjust to them, without requiring a nervous system. On a molecular level, auxins have an aromatic ring and a carboxylic acid group (Taiz and Zeiger, 1998).

The most important member of the auxin family is indole-3-acetic acid (IAA). It generates the majority of auxin effects in intact plants, and is the most potent native auxin. However, molecules of IAA are chemically labile in aqueous solution, so IAA is not used commercially as a plant growth regulator.

• Naturally-occurring auxins include 4-chloro-indoleacetic acid, phenylacetic acid (PAA) and indole-3-butyric acid (IBA).
• Synthetic auxin analogs include 1-naphthaleneacetic acid (NAA), 2,4-dichlorophenoxyacetic acid (2,4-D), and others.

Auxins are often used to promote initiation of adventitious roots and are the active ingredient of the commercial preparations used in horticulture to root stem cuttings. They can also be used to promote uniform flowering, to promote fruit set, and to prevent premature fruit drop.

Used in high doses, auxin stimulates the production of ethylene. Excess ethylene can inhibit elongation growth, cause leaves to fall (leaf abscission), and even kill the plant. Some synthetic auxins such as 2,4-D and 2,4,5-trichlorophenoxyacetic acid (2,4,5-T) have been used as herbicides. Broad-leaf plants (dicots) such as dandelions are much more susceptible to auxins than narrow-leaf plants (monocots) like grass and cereal crops. These synthetic auxins were the active agents in Agent Orange, a defoliant used extensively by American forces in the Vietnam War.

Hormonal activity

Auxins coordinate development at all levels in plants, from the cellular level to organs and ultimately the whole plant.

Molecular mechanisms

Auxins directly stimulate or inhibit the expression of specific genes. Auxin induces transcription by targeting for degradation members of the Aux/IAA family of transcriptional repressor proteins, The degradation of the Aux/IAAs leads to the derepression of Auxin Respose Factors ARF-mediated transcription. Aux/IAAs are targeted for degradation by ubiquitination, catalysed by an SCF-type ubiquitin-protein ligase.

In 2005, it was demonstrated that the F-box protein TIR1, which is part of the ubiquitin ligase complex SCFTIR1, is an auxin receptor. Upon auxin binding TIR1 recruits specific transcriptional repressors (the Aux/IAA repressors) for ubiquitination by the SCF complex. This marking process leads to the degradation of the repressors by the proteasome, alleviating repression and leading to specific gene expression in response to auxins (reviewed in ).

Another protein called ABP1 (Auxin Binding Protein 1) is a putative receptor, but its role is unclear. Electrophysiological experiments with protoplasts and anti-ABP1 antibodies suggest that ABP1 may have a function at the plasma membrane.

On a cellular level

On the cellular level, auxin is essential for cell growth, affecting both cell division and cellular expansion. Depending on the specific tissue, auxin may promote axial elongation (as in shoots), lateral expansion (as in root swelling), or isodiametric expansion (as in fruit growth). In some cases (coleoptile growth) auxin-promoted cellular expansion occurs in the absence of cell division. In other cases, auxin-promoted cell division and cell expansion may be closely sequenced within the same tissue (root initiation, fruit growth). In a living plant it appears that auxins and other plant hormones nearly always interact to determine patterns of plant development.

According to the acid growth hypothesis for auxin action, auxins may directly stimulate the early phases of cell elongation by causing responsive cells to actively transport hydrogen ions out of the cell, thus lowering the pH around cells. This acidification of the cell wall region activates wall-loosening proteins known as expansins, which allow slippage of cellulose microfibrils in the cell wall, making the cell wall less rigid. When the cell wall is loosened by the action of auxins, this now-less-rigid wall is expanded by cell turgor pressure, which presses against the cell wall.

However, the acid growth hypothesis does not by itself account for the increased synthesis and transport of cell wall precursors and secretory activity in the Golgi system that accompany and sustain auxin-promoted cell expansion.

Organ patterns

Growth and division of plant cells together result in growth of tissue, and specific tissue growth contributes to the development of plant organs. Growth of cells contributes to the plant's size, but uneven localized growth produces bending, turning and directionalization of organs- for example, stems turning toward light sources (phototropism), roots growing in response to gravity (gravitropism), and other tropisms.

Organization of the plant

As auxins contribute to organ shaping, they are also fundamentally required for proper development of the plant itself. Without hormonal regulation and organization, plants would be merely proliferating heaps of similar cells. Auxin employment begins in the embryo of the plant, where directional distribution of auxin ushers in subsequent growth and development of primary growth poles, then forms buds of future organs. Throughout the plant's life, auxin helps the plant maintain the polarity of growth and recognize where it has its branches (or any organ) connected.

An important principle of plant organization based upon auxin distribution is apical dominance, which means that the auxin produced by the apical bud (or growing tip) diffuses downwards and inhibits the development of ulterior lateral bud growth, which would otherwise compete with the apical tip for light and nutrients. Removing the apical tip and its suppressive hormone allows the lower dormant lateral buds to develop, and the buds between the leaf stalk and stem produce new shoots which compete to become the lead growth. This behavior is used in pruning by horticulturists.

Uneven distribution of auxin: To cause growth in the required domains, it is necessary that auxins be active preferentially in them. Auxins are not synthesized everywhere, but each cell retains the potential ability to do so, and only under specific conditions will auxin synthesis be activated. For that purpose, not only do auxins have to be translocated toward those sites where they are needed but there has to be an established mechanism to detect those sites. Translocation is driven throughout the plant body primarily from peaks of shoots to peaks of roots. For long distances, relocation occurs via the stream of fluid in phloem vessels, but, for short-distance transport, a unique system of coordinated polar transport directly from cell to cell is exploited. This process of polar auxin transport is directional and very strictly regulated. It is based in uneven distribution of auxin efflux carriers on the plasma membrane, which send auxins in the proper direction.

A 2006 study showed plant-specific pin-formed (PIN) proteins are vital in transporting auxin. PINs also regulate auxin efflux from mammalian and yeast cells.

Locations

• In shoot (and root) meristematic tissue
• In young leaves
• In mature leaves in very tiny amounts
• In mature root cells in even smaller amounts
• Transported throughout the plant more prominently downward from the shoot apices

Effects

The plant hormone stimulates cell elongation. It stimulates the Wall Loosening Factors, for example, elastins, to loosen the cell walls. If gibberellins are also present, the effect is stronger. It also stimulates cell division if cytokinins are present. When auxin and cytokinin are applied to callus, rooting can be generated if the auxin concentration is higher than cytokinin concentration while xylem tissues can be generated when the auxin concentration is equal to the cytokinins.

It participates in phototropism, geotropism, hydrotropism and other developmental changes. The uneven distribution of auxin, due to environmental cues (for example, unidirectional light and gravity force), results in uneven plant tissue growth.

It also induces sugar and mineral accumulation at the site of application.

Wounding response

It induces formation and organization of phloem and xylem. When the plant is wounded, the auxin may induce the Cell differentiation and regeneration of the vascular tissues.

Root growth and development

Auxin induces new root formation by breaking root apical dominance induced by cytokinins. In horticulture, auxins, especially NAA and IBA, are commonly applied to stimulate root growth when taking cuttings of plants. However, high concentrations of auxin inhibit root elongation and instead enhance adventitious root formation. Removal of the root tip can lead to inhibition of secondary root formation.

Apical dominance

It induces shoot apical dominance; the axillary buds are inhibited by auxin. When the apex of the plant is removed, the inhibitory effect is removed and the growth of lateral buds is enhanced as a high concentration of auxin directly stimulates ethylene synthesis in lateral buds causes inhibition of its growth and potentiation of apical dominance.

Ethylene biosynthesis

In low concentrations, auxin can inhibit ethylene formation and transport of precursor in plants; however, high concentrations of auxin can induce the synthesis of ethylene. Therefore, the high concentration can induce femaleness of flowers in some species.

It inhibits abscission prior to formation of abscission layer and thus inhibits senescence of leaves.

Fruit growth

Auxin delays fruit senescence.

It is required for fruit growth. When seeds are removed from strawberries, fruit growth is stopped; exogenous auxin stimulates the growth in seed removed fruits. For fruit with unfertilized seeds, exogenous auxin results in parthenocarpy ("virgin-fruit" growth).

Flowering

Auxin plays a minor role in the initiation of flowering. It can delay the senescence of flowers in low concentrations.

Herbicide manufacture

The defoliant Agent Orange was a mix of 2,4-D and 2,4,5-T. The compound 2,4-D is still in use and is thought to be safe, but 2,4,5-T was more or less banned by the EPA in 1979. The dioxin TCDD is an unavoidable contaminant produced in the manufacture of 2,4,5-T. As a result of the integral dioxin contamination, 2,4,5-T has been implicated in leukaemia, miscarriages, birth defects, liver damage, and other diseases. Agent Orange was sprayed in Vietnam as a defoliant to deny ground cover to the Vietnamese army.

See also

• Herbicide
• Pruning fruit trees
• Fusicoccin

References

• Plant Physiology Online - Chapter 19: Auxin: The Growth Hormone
• Plant Physiology

Taiz, L. & Zeiger, E. (1998). Plant Physiology. 2nd edition. Massachusetts: Sinauer Associates, Inc. 792 p.