Auxin
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Auxins (plural of auxin ) are a class of
plant hormone Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pa ...
s (or plant-growth regulators) with some
morphogen A morphogen is a substance whose non-uniform distribution governs the pattern of tissue development in the process of morphogenesis or pattern formation, one of the core processes of developmental biology, establishing positions of the various ...
-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 Frits Warmolt Went (May 18, 1903 – May 1, 1990) was a Dutch biologist whose 1928 experiment demonstrated the existence of auxin in plants. Went's father was the prominent Dutch botanist F.A.F.C. Went. After graduating from the University of ...
first described auxins and their role in plant growth in the 1920s. Kenneth V. Thimann became the first to isolate one of these
phytohormones Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pa ...
and to determine its chemical structure as
indole-3-acetic acid Indole-3-acetic acid (IAA, 3-IAA) is the most common naturally occurring plant hormone of the auxin class. It is the best known of the auxins, and has been the subject of extensive studies by plant physiologists. IAA is a derivative of indole, con ...
(IAA). Went and Thimann co-authored a book on plant hormones, ''Phytohormones'', in 1937.


Overview

Auxins were the first of the major
plant hormone Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pa ...
s to be discovered. They derive their name from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
word αυξειν (''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 A pattern is a regularity in the world, in human-made design, or in abstract ideas. As such, the elements of a pattern repeat in a predictable manner. A geometric pattern is a kind of pattern formed of geometric shapes and typically repeated l ...
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 A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
or
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s). It is achieved through very complex and well-coordinated
active Active may refer to: Music * ''Active'' (album), a 1992 album by Casiopea * Active Records, a record label Ships * ''Active'' (ship), several commercial ships by that name * HMS ''Active'', the name of various ships of the British Royal ...
transport of auxin molecules from cell to cell throughout the plant body — by the so-called
polar auxin transport Polar auxin transport is the regulated transport of the plant hormone auxin in plants. It is an active process, the hormone is transported in cell-to-cell manner and one of the main features of the transport is its asymmetry and directionality ( p ...
. Thus, a plant can (as a whole) react to external conditions and adjust to them, without requiring a
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...
. Auxins typically act in concert with, or in opposition to, other plant hormones. For example, the ratio of auxin to
cytokinin Cytokinins (CK) are a class of plant hormones that promote cell division, or cytokinesis, in plant roots and shoots. They are involved primarily in cell growth and differentiation, but also affect apical dominance, axillary bud growth, and le ...
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 In organic chemistry, a carboxylic acid is an organic acid that contains a carboxyl group () attached to an R-group. The general formula of a carboxylic acid is or , with R referring to the alkyl, alkenyl, aryl, or other group. Carboxylic ...
group. The most important member of the auxin family is
indole-3-acetic acid Indole-3-acetic acid (IAA, 3-IAA) is the most common naturally occurring plant hormone of the auxin class. It is the best known of the auxins, and has been the subject of extensive studies by plant physiologists. IAA is a derivative of indole, con ...
(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 Conjugation or conjugate may refer to: Linguistics * Grammatical conjugation, the modification of a verb from its basic form * Emotive conjugation or Russell's conjugation, the use of loaded language Mathematics * Complex conjugation, the chang ...
to degradation of its molecules, always according to the requirements of the situation. * Five naturally occurring (endogenous) auxins in plants include indole-3-acetic acid, 4-chloroindole-3-acetic acid, phenylacetic acid,
indole-3-butyric acid Indole-3-butyric acid (1''H''-indole-3-butanoic acid, IBA) is a white to light-yellow crystalline solid, with the molecular formula C12H13NO2. It melts at 125 °C in atmospheric pressure and decomposes before boiling. IBA is a plant hormone ...
, and
indole-3-propionic acid 3-Indolepropionic acid (IPA), or indole-3-propionic acid, is a potent neuroprotective antioxidant, plant auxin, and natural product in humans that is being studied for therapeutic use in Alzheimer's disease. It is endogenously produced by h ...
. However, most of the knowledge described so far in auxin biology and as described in the sections which follow, apply basically to IAA; the other three endogenous auxins seems to have marginal importance for intact plants in natural environments. Alongside endogenous auxins, scientists and manufacturers have developed many synthetic compounds with auxinic activity. * Synthetic auxin analogs include 1-naphthaleneacetic acid, 2,4-dichlorophenoxyacetic acid (2,4-D), and many others. Some synthetic auxins, such as 2,4-D and 2,4,5-trichlorophenoxyacetic acid (2,4,5-T), are sold as
herbicide Herbicides (, ), also commonly known as weedkillers, are substances used to control undesired plants, also known as weeds.EPA. February 201Pesticides Industry. Sales and Usage 2006 and 2007: Market Estimates. Summary in press releasMain page fo ...
s. Broad-leaf plants ( dicots), such as
dandelion ''Taraxacum'' () is a large genus of flowering plants in the family Asteraceae, which consists of species commonly known as dandelions. The scientific and hobby study of the genus is known as taraxacology. The genus is native to Eurasia and Nor ...
s, are much more susceptible to auxins than narrow-leaf plants (
monocots Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of t ...
) such as
grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns an ...
es and
cereal A cereal is any Poaceae, grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, Cereal germ, germ, and bran. Cereal Grain, grain crops are grown in greater quantit ...
crops, making these synthetic auxins valuable as herbicides.


Discovery of auxin


Charles Darwin

In 1881,
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
and his son
Francis Francis may refer to: People *Pope Francis, the head of the Catholic Church and sovereign of the Vatican City State and Bishop of Rome *Francis (given name), including a list of people and fictional characters *Francis (surname) Places * Rural M ...
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 The hypocotyl (short for "hypocotyledonous stem", meaning "below seed leaf") is the stem of a germinating seedling, found below the cotyledons (seed leaves) and above the radicle ( root). Eudicots As the plant embryo grows at germination, it se ...
. 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.


Peter Boysen Jensen

In 1910,
Danish Danish may refer to: * Something of, from, or related to the country of Denmark People * A national or citizen of Denmark, also called a "Dane," see Demographics of Denmark * Culture of Denmark * Danish people or Danes, people with a Danish a ...
scientist demonstrated that the
phototropic Phototropism is the growth of an organism in response to a light stimulus. Phototropism is most often observed in plants, but can also occur in other organisms such as fungi. The cells on the plant that are farthest from the light contain a hor ...
stimulus A stimulus is something that causes a physiological response. It may refer to: *Stimulation **Stimulus (physiology), something external that influences an activity **Stimulus (psychology), a concept in behaviorism and perception *Stimulus (economi ...
in the
oat The oat (''Avena sativa''), sometimes called the common oat, is a species of cereal grain grown for its seed, which is known by the same name (usually in the plural, unlike other cereals and pseudocereals). While oats are suitable for human con ...
coleoptile Coleoptile is the pointed protective sheath covering the emerging shoot in monocotyledons such as grasses in which few leaf primordia and shoot apex of monocot embryo remain enclosed. The coleoptile protects the first leaf as well as the growing ...
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 Micas ( ) are a group of silicate minerals whose outstanding physical characteristic is that individual mica crystals can easily be split into extremely thin elastic plates. This characteristic is described as perfect basal cleavage. Mica is ...
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
tropism A tropism is a biological phenomenon, indicating growth or turning movement of a biological organism, usually a plant, in response to an environmental stimulus. In tropisms, this response is dependent on the direction of the stimulus (as oppos ...
s.


Frits Went

In 1928, the Dutch botanist
Frits Warmolt Went Frits Warmolt Went (May 18, 1903 – May 1, 1990) was a Dutch biologist whose 1928 experiment demonstrated the existence of auxin in plants. Went's father was the prominent Dutch botanist F.A.F.C. Went. After graduating from the University of ...
showed that a chemical messenger diffuses from coleoptile tips. Went's experiment identified how a growth promoting chemical causes a coleoptile to grow towards the light. Went cut the tips of the coleoptiles and placed them in the dark, putting a few tips on agar blocks that he predicted would absorb the growth-promoting chemical. On control coleoptiles, he placed a block that lacked the chemical. On others, he placed blocks containing the chemical, either centered on top of the coleoptile to distribute the chemical evenly or offset to increase the concentration on one side. When the growth-promoting chemical was distributed evenly the coleoptile grew straight. If the chemical was distributed unevenly, the coleoptile curved away from the side with the cube, as if growing towards the light, even though it was grown in the dark. Went later proposed that the messenger substance is a growth-promoting hormone, which he named auxin, that becomes asymmetrically distributed in the bending region. Went concluded that auxin is at a higher concentration on the shaded side, promoting cell elongation, which results in coleoptiles bending towards the light.


Hormonal activity

Auxins help development at all levels in plants, from the
cellular Cellular may refer to: *Cellular automaton, a model in discrete mathematics * Cell biology, the evaluation of cells work and more * ''Cellular'' (film), a 2004 movie *Cellular frequencies, assigned to networks operating in cellular RF bands *Cell ...
level, through organs, and ultimately to the whole plant.


Molecular mechanisms

When a plant cell comes into contact with auxin, it causes dramatic changes in
gene expression Gene expression is the process by which information from a gene is used in the synthesis of a functional gene product that enables it to produce end products, protein or non-coding RNA, and ultimately affect a phenotype, as the final effect. The ...
, with many genes up- or down-regulated. The precise mechanisms by which this occurs are still an area of active research, but there is now a general consensus on at least two auxin signalling pathways.


Perception

The most well-characterized auxin receptors are the TIR1/ AFB family of
F-box protein F-box proteins are proteins containing at least one F-box domain. The first identified F-box protein is one of three components of the SCF complex, which mediates ubiquitination of proteins targeted for degradation by the 26S proteasome. Core c ...
s. F-box proteins target other proteins for degradation via the ubiquitin degradation pathway. When TIR1/ AFB proteins bind to auxin, the auxin acts as a 'molecular glue' that allows these proteins to then bind to their targets (see below). Another auxin-binding protein, ABP1 is now often regarded as an auxin receptor (at the
apoplast Inside a plant, the apoplast can mean the space outside of cell membranes, where material can diffuse freely; that is, the extracellular spaces. ''Apoplast '' can also refer especially to the continuum of cell walls of adjacent cells; fluid and ...
), but it is generally considered to have a much more minor role than the TIR1/AFB signaling pathway, and much less is known about ABP1 signaling.


Aux/IAA and ARF signalling modules

Auxin response factors (ARFs) are a large group of
transcription factor In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The fu ...
s that act in auxin signaling. In the absence of auxin, ARFs bind to a class of repressors known as Aux/IAAs. Aux/IAA suppress the ability of ARFs to enhance gene transcription. Additionally, the binding of Aux/IAA to ARFs brings Aux/IAA into contact with the promoters of auxin-regulated genes. When at these promoters, Aux/IAA repress the expression of these genes through recruiting other factors to make modifications to the DNA structure. The binding of auxin to TIR1/AFBs allows them to bind to Aux/IAAs. When bound by TIR1/AFBs, Aux/IAAs are marked for degradation. The degradation of Aux/IAA frees ARF proteins, which are then able to activate or repress genes at whose promoters they are bound. The large number of Aux/IAA and ARF binding pairs possible, and their different distributions between cell types and across developmental age are thought to account for the astonishingly diverse responses that auxin produces. In June 2018, it was demonstrated that plant tissues can respond to auxin in a TIR1-dependent manner extremely quickly (probably too quickly to be explained by changes in gene expression). This has led some scientists to suggest that there is an as yet unidentified TIR1-dependent auxin-signalling pathway that differs from the well-known transcriptional response.


On a cellular level

On the cellular level, auxin is essential for
cell growth Cell growth refers to an increase in the total mass of a cell, including both cytoplasmic, nuclear and organelle volume. Cell growth occurs when the overall rate of cellular biosynthesis (production of biomolecules or anabolism) is greater than ...
, affecting both
cell division Cell division is the process by which a parent cell (biology), cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukar ...
and cellular expansion. Auxin concentration level, together with other local factors, contributes to cell differentiation and specification of the cell fate. Depending on the specific tissue, auxin may promote axial elongation (as in shoots), lateral expansion (as in root swelling), or iso-diametric 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, auxins and other plant hormones nearly always appear to interact to determine patterns of plant development.


Organ patterns

Growth and division of plant cells together result in the growth of tissue, and specific tissue growth contributes to the development of plant organs. Growth of cells contributes to the plant's size, unevenly localized growth produces bending, turning and directionalization of organs- for example, stems turning toward light sources ( phototropism), roots growing in response to gravity (
gravitropism Gravitropism (also known as geotropism) is a coordinated process of differential growth by a plant in response to gravity pulling on it. It also occurs in fungi. Gravity can be either "artificial gravity" or natural gravity. It is a general featu ...
), and other
tropism A tropism is a biological phenomenon, indicating growth or turning movement of a biological organism, usually a plant, in response to an environmental stimulus. In tropisms, this response is dependent on the direction of the stimulus (as oppos ...
s originated because cells on one side grow faster than the cells on the other side of the organ. So, precise control of auxin distribution between different cells has paramount importance to the resulting form of plant growth and organization.


Auxin transport and the uneven distribution of auxin

To cause growth in the required domains, auxins must of necessity be active preferentially in them. Local auxin maxima can be formed by active biosynthesis in certain cells of tissues, for example via tryptophan-dependent pathways, but auxins are not synthesized in all cells (even if cells retain the potential ability to do so, only under specific conditions will auxin synthesis be activated in them). For that purpose, auxins have to be not only translocated toward those sites where they are needed but also they must have an established mechanism to detect those sites. Translocation is driven throughout the plant body, primarily from peaks of shoots to peaks of roots (from up to down). For long distances, relocation occurs via the stream of fluid in
phloem Phloem (, ) is the living biological tissue, tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This tran ...
vessels, but, for short-distance transport, a unique system of coordinated polar transport directly from cell to cell is exploited. This short-distance, active transport exhibits some
morphogen A morphogen is a substance whose non-uniform distribution governs the pattern of tissue development in the process of morphogenesis or pattern formation, one of the core processes of developmental biology, establishing positions of the various ...
etic properties. This process,
polar auxin transport Polar auxin transport is the regulated transport of the plant hormone auxin in plants. It is an active process, the hormone is transported in cell-to-cell manner and one of the main features of the transport is its asymmetry and directionality ( p ...
, is directional, very strictly regulated, and based in uneven distribution of auxin efflux carriers on the plasma membrane, which send auxins in the proper direction. While PIN-FORMED (PIN) proteins are vital in transporting auxin in a polar manner, the family of AUXIN1/LIKE-AUX1 (AUX/LAX) genes encodes for non-polar auxin influx carriers. The regulation of PIN protein localisation in a cell determines the direction of auxin transport from cell, and concentrated effort of many cells creates peaks of auxin, or auxin maxima (regions having cells with higher auxin – a maximum). Proper and timely auxin maxima within developing roots and shoots are necessary to organise the development of the organ. PINs are regulated by multiple pathways, at both the transcriptional and the post-translational levels. PIN proteins can be phosphorylated by PINOID, which determines their apicobasal polarity and thereby the directionality of auxin fluxes. In addition, other AGC kinases, such as D6PK, phosphorylate and activate PIN transporters. AGC kinases, including PINOID and D6PK, target to the plasma membrane via binding to phospholipids. Upstream of D6PK, 3'-phosphoinositide dependent protein kinase 1 (PDK1) acts as a master regulator. PDK1 phosphorylates and activates D6PK at the basal side of plasma membrane, executing the activity of PIN-mediated polar auxin transport and subsequent plant development. Surrounding auxin maxima are cells with low auxin troughs, or auxin minima. For example, in the ''
Arabidopsis ''Arabidopsis'' (rockcress) is a genus in the family Brassicaceae. They are small flowering plants related to cabbage and mustard. This genus is of great interest since it contains thale cress (''Arabidopsis thaliana''), one of the model organi ...
'' fruit, auxin minima have been shown to be important for its tissue development. Auxin has a significant effect on spatial and temporal gene expressions during the growth of apical meristems. These interactions depend both on the concentration of Auxin as well as the spatial orientation during primordial positioning. Auxin relies on PIN1 which works as an auxin efflux carrier. PIN1 positioning upon membranes determines the directional flow of the hormone from higher to lower concentrations. Initiation of primordia in apical meristems is correlated to heightened auxin levels. Genes required to specify the identity of cells arrange and express based on levels of auxin. STM (SHOOT MERISTEMLESS), which helps maintain undifferentiated cells, is down-regulated in the presence of auxin. This allows growing cells to differentiate into various plant tissues. The CUC (CUP-SHAPED COTYLEDON) genes set the boundaries for growing tissues and promote growth. They are upregulated via auxin influx. Experiments making use of GFP (GREEN FLUORESCENCE PROTEIN) visualization in Arabidopsis have supported these claims.


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 the directional distribution of auxin ushers in subsequent growth and development of primary growth poles, then forms buds of future organs. Next, it helps to coordinate proper development of the arising organs, such as roots, cotyledons, and leaves and mediates long-distance signals between them, contributing so to the overall architecture of the plant. Throughout the plant's life, auxin helps the plant maintain the polarity of growth, and actually "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 the auxin produced by the apical bud (or growing tip) diffuses (and is transported) 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 suppressively acting auxin 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. The process is actually quite complex because auxin transported downwards from the lead shoot tip has to interact with several other plant hormones (such as
strigolactone Strigolactones are a group of chemical compounds produced by a plant's roots. Due to their mechanism of action, these molecules have been classified as plant hormones or phytohormones. So far, strigolactones have been identified to be responsible fo ...
s or
cytokinins Cytokinins (CK) are a class of plant hormones that promote cell division, or cytokinesis, in plant roots and shoots. They are involved primarily in cell growth and differentiation, but also affect apical dominance, axillary bud growth, and le ...
) in the process on various positions along the growth axis in plant body to achieve this phenomenon. This plant behavior is used in
pruning Pruning is a horticultural, arboricultural, and silvicultural practice involving the selective removal of certain parts of a plant, such as branches, buds, or roots. The practice entails the ''targeted'' removal of diseased, damaged, dead, ...
by horticulturists. Finally, the sum of auxin arriving from stems to roots influences the degree of root growth. If shoot tips are removed, the plant does not react just by the outgrowth of lateral buds — which are supposed to replace to original lead. It also follows that smaller amount of auxin arriving at the roots results in slower growth of roots and the nutrients are subsequently in higher degree invested in the upper part of the plant, which hence starts to grow faster.


Effects

Auxin participates in phototropism,
geotropism Gravitropism (also known as geotropism) is a coordinated process of differential growth by a plant in response to gravity pulling on it. It also occurs in fungi. Gravity can be either "artificial gravity" or natural gravity. It is a general featu ...
,
hydrotropism Hydrotropism (hydro- "water"; tropism "involuntary orientation by an organism, that involves turning or curving as a positive or negative response to a stimulus") is a plant's growth response in which the direction of growth is determined by a sti ...
and other developmental changes. The uneven distribution of auxin, due to environmental cues, such as unidirectional light or gravity force, results in uneven plant tissue growth, and generally, auxin governs the form and shape of the plant body, direction and strength of growth of all organs, and their mutual interaction. When the cells grow larger, their volume increases as the intracellular solute concentration increases with water moving into the cells from extracellular fluid. This auxin-stimulated intake of water causes turgor pressure on the cell walls, causing the plant to bend. Auxin stimulates cell elongation by stimulating wall-loosening factors, such as
expansin Expansins are a family of closely related nonenzymatic proteins found in the plant cell wall, with important roles in plant cell growth, fruit softening, abscission, emergence of root hairs, pollen tube invasion of the stigma and style, meristem fun ...
s, to loosen
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
s. The effect is stronger if gibberellins are also present. Auxin also stimulates cell division if
cytokinins Cytokinins (CK) are a class of plant hormones that promote cell division, or cytokinesis, in plant roots and shoots. They are involved primarily in cell growth and differentiation, but also affect apical dominance, axillary bud growth, and le ...
are present. When auxin and cytokinin are applied to
callus A callus is an area of thickened and sometimes hardened skin that forms as a response to repeated friction, pressure, or other irritation. Since repeated contact is required, calluses are most often found on the feet and hands, but they may o ...
, rooting can be generated with higher auxin to cytokinin ratios, shoot growth is induced by lower auxin to cytokinin ratios, and a callus is formed with intermediate ratios, with the exact threshold ratios depending on the species and the original tissue. Auxin also induces sugar and mineral accumulation at the site of application.


Wound response

Auxin induces the formation and organization of
phloem Phloem (, ) is the living biological tissue, tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This tran ...
and
xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
. When the plant is wounded, the auxin may induce the cell differentiation and regeneration of the vascular tissues.


Root growth and development

Auxins promote root initiation. Auxin induces both growth of pre-existing roots and root branching (lateral root initiation), and also adventitious root formation. As more native auxin is transported down the stem to the roots, the overall development of the roots is stimulated. If the source of auxin is removed, such as by trimming the tips of stems, the roots are less stimulated accordingly, and growth of stem is supported instead. In horticulture, auxins, especially
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and IBA, are commonly applied to stimulate root initiation when rooting 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

Auxin induces shoot apical dominance; the
axillary bud The axillary bud (or lateral bud) is an embryonic or organogenic shoot located in the axil of a leaf. Each bud has the potential to form shoots, and may be specialized in producing either vegetative shoots (stems and branches) or reproductive sh ...
s are inhibited by auxin, as a high concentration of auxin directly stimulates
ethylene Ethylene (IUPAC name: ethene) is a hydrocarbon which has the formula or . It is a colourless, flammable gas with a faint "sweet and musky" odour when pure. It is the simplest alkene (a hydrocarbon with carbon-carbon double bonds). Ethylene i ...
synthesis in axillary buds, causing inhibition of their growth and potentiation of apical dominance. When the apex of the plant is removed, the inhibitory effect is removed and the growth of lateral buds is enhanced. This is called decapitation, usually performed in tea plantations and hedge-making. Auxin is sent to the part of the plant facing away from the light, where it promotes cell elongation, thus causing the plant to bend towards the light.


Fruit growth and development

Auxin is required for fruit growth and development and delays fruit
senescence Senescence () or biological aging is the gradual deterioration of functional characteristics in living organisms. The word ''senescence'' can refer to either cellular senescence or to senescence of the whole organism. Organismal senescence inv ...
. When seeds are removed from strawberries, fruit growth is stopped; exogenous auxin stimulates the growth in fruits with seeds removed. For fruit with unfertilized seeds, exogenous auxin results in
parthenocarpy In botany and horticulture, parthenocarpy is the natural or artificially induced production of fruit without fertilisation of ovules, which makes the fruit seedless. Stenospermocarpy may also produce apparently seedless fruit, but the seeds are ...
("virgin-fruit" growth). Fruits form abnormal morphologies when auxin transport is disturbed. In ''
Arabidopsis ''Arabidopsis'' (rockcress) is a genus in the family Brassicaceae. They are small flowering plants related to cabbage and mustard. This genus is of great interest since it contains thale cress (''Arabidopsis thaliana''), one of the model organi ...
'' fruits, auxin controls the release of seeds from the fruit (pod). The valve margins are a specialised tissue in pods that regulates when pod will open (dehiscence). Auxin must be removed from the valve margin cells to allow the valve margins to form. This process requires modification of the auxin transporters (PIN proteins). The evolutionary transition from
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
to
triploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei ( eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contain ...
endosperm The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid (meaning three chromosome sets per nucleus) in most species, which may be auxin-driven. It surrounds the embryo and ...
s - and the production of antipodal cells - may have occurred due to a shift in
gametophyte A gametophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has on ...
development which produced a new interaction with an auxin-dependent mechanism originating in the earliest
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s.


Flowering

Auxin plays also a minor role in the initiation of flowering and development of reproductive organs. In low concentrations, it can delay the
senescence Senescence () or biological aging is the gradual deterioration of functional characteristics in living organisms. The word ''senescence'' can refer to either cellular senescence or to senescence of the whole organism. Organismal senescence inv ...
of flowers. A number of plant mutants have been described that affect flowering and have deficiencies in either auxin synthesis or transport. In maize, one example is bif2 barren inflorescence2.


Ethylene biosynthesis

In low concentrations, auxin can inhibit ethylene formation and transport of precursor in plants; however, high concentrations can induce the synthesis of ethylene. Therefore, the high concentration can induce femaleness of flowers in some species. Auxin inhibits
abscission Abscission () is the shedding of various parts of an organism, such as a plant dropping a leaf, fruit, flower, or seed. In zoology, abscission is the intentional shedding of a body part, such as the shedding of a claw, husk, or the autotomy of a ...
prior to the formation of the abscission layer, and thus inhibits senescence of leaves.


Synthetic auxins

In the course of research on auxin biology, many compounds with noticeable auxin activity were synthesized. Many of them had been found to have economical potential for human-controlled growth and development of plants in agronomy. Auxins are toxic to plants in large concentrations; they are most toxic to
dicot The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, t ...
s and less so to
monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...
s. Because of this property, synthetic auxin herbicides, including 2,4-dichlorophenoxyacetic acid (2,4-D) and 2,4,5-trichlorophenoxyacetic acid (2,4,5-T), have been developed and used for
weed A weed is a plant considered undesirable in a particular situation, "a plant in the wrong place", or a plant growing where it is not wanted.Harlan, J. R., & deWet, J. M. (1965). Some thoughts about weeds. ''Economic botany'', ''19''(1), 16-24. ...
control. However, some exogenously synthesized auxins, especially 1-naphthaleneacetic acid (NAA) and
indole-3-butyric acid Indole-3-butyric acid (1''H''-indole-3-butanoic acid, IBA) is a white to light-yellow crystalline solid, with the molecular formula C12H13NO2. It melts at 125 °C in atmospheric pressure and decomposes before boiling. IBA is a plant hormone ...
(IBA), are also commonly applied to stimulate root growth when taking cuttings of plants or for different agricultural purposes such as the prevention of fruit drop in
orchard An orchard is an intentional plantation of trees or shrubs that is maintained for food production. Orchards comprise fruit- or nut-producing trees which are generally grown for commercial production. Orchards are also sometimes a feature of larg ...
s. Used in high doses, auxin stimulates the production of
ethylene Ethylene (IUPAC name: ethene) is a hydrocarbon which has the formula or . It is a colourless, flammable gas with a faint "sweet and musky" odour when pure. It is the simplest alkene (a hydrocarbon with carbon-carbon double bonds). Ethylene i ...
, also a native plant hormone. Excess ethylene can inhibit elongation growth, cause
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
to fall (
abscission Abscission () is the shedding of various parts of an organism, such as a plant dropping a leaf, fruit, flower, or seed. In zoology, abscission is the intentional shedding of a body part, such as the shedding of a claw, husk, or the autotomy of a ...
), and even kill the plant. Some synthetic auxins, such as 2,4-D and 2,4,5-T are marketed also as
herbicide Herbicides (, ), also commonly known as weedkillers, are substances used to control undesired plants, also known as weeds.EPA. February 201Pesticides Industry. Sales and Usage 2006 and 2007: Market Estimates. Summary in press releasMain page fo ...
s. Dicots, such as
dandelion ''Taraxacum'' () is a large genus of flowering plants in the family Asteraceae, which consists of species commonly known as dandelions. The scientific and hobby study of the genus is known as taraxacology. The genus is native to Eurasia and Nor ...
s, are much more susceptible to auxins than
monocots Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of t ...
, such as
grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns an ...
es and
cereal A cereal is any Poaceae, grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, Cereal germ, germ, and bran. Cereal Grain, grain crops are grown in greater quantit ...
crops. So these synthetic auxins are valuable as synthetic herbicides. 2,4-D was the first widely used herbicide, and it is still in use. 2,4-D was first commercialized by the
Sherwin-Williams Sherwin-Williams Company is an American Cleveland, Ohio–based company in the paint and coating manufacturing industry. The company primarily engages in the manufacture, distribution, and sale of paints, coatings, floorcoverings, and related p ...
company and saw use in the late 1940s. It is easy and inexpensive to manufacture.
Triclopyr Triclopyr (3,5,6-trichloro-2-pyridinyloxyacetic acid) is an organic compound in the pyridine group that is used as a systemic foliar herbicide and fungicide. Uses Triclopyr is used to control broadleaf weeds while leaving grasses and conifers ...
(3,5,6-TPA), while known as an herbicide, has also been shown to increase the size of fruit in plants. At increased concentrations, the hormone can be lethal. Dosing down to the correct concentration has been shown to alter photosynthetic pathways. This hindrance to the plant causes a response that increases carbohydrate production, leading to larger fruit.


Herbicide manufacture

Synthetic auxins are used as a kind of herbicide and overdosing of auxins will interrupt plants' growth and lead to their death. The
defoliant A defoliant is any herbicidal chemical sprayed or dusted on plants to cause their leaves to fall off. Defoliants are widely used for the selective removal of weeds in managing croplands and lawns. Worldwide use of defoliants, along with the ...
Agent Orange, used extensively by British forces in the Malayan Emergency and American forces in the
Vietnam War The Vietnam War (also known by #Names, other names) was a conflict in Vietnam, Laos, and Cambodia from 1 November 1955 to the fall of Saigon on 30 April 1975. It was the second of the Indochina Wars and was officially fought between North Vie ...
, 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
U.S. Environmental Protection Agency The Environmental Protection Agency (EPA) is an Independent agencies of the United States government, independent executive agency of the United States federal government tasked with environmental protection matters. President Richard Nixon pro ...
in 1979. The
dioxin Dioxin may refer to: * 1,2-Dioxin or 1,4-Dioxin, two unsaturated heterocyclic 6-membered rings where two carbon atoms have been replaced by oxygen atoms, giving the molecular formula C4H4O2 *Dibenzo-1,4-dioxin, the parent compound also known as ...
TCDD TCDD may refer to any of the following: * Turkish State Railways The State Railways of the Republic of Turkey ( tr, Türkiye Cumhuriyeti Devlet Demiryolları), abbreviated as TCDD, is a government-owned national railway company responsible with ...
is an unavoidable contaminant produced in the manufacture of 2,4,5-T. As a result of the integral dioxin contamination, the use of 2,4,5-T products has been implicated in
leukemia Leukemia ( also spelled leukaemia and pronounced ) is a group of blood cancers that usually begin in the bone marrow and result in high numbers of abnormal blood cells. These blood cells are not fully developed and are called ''blasts'' or ' ...
,
miscarriage Miscarriage, also known in medical terms as a spontaneous abortion and pregnancy loss, is the death of an embryo or fetus before it is able to survive independently. Miscarriage before 6 weeks of gestation is defined by ESHRE as biochemical lo ...
s,
birth defect A birth defect, also known as a congenital disorder, is an abnormal condition that is present at birth regardless of its cause. Birth defects may result in disabilities that may be physical, intellectual, or developmental. The disabilities ca ...
s,
liver The liver is a major Organ (anatomy), organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the Protein biosynthesis, synthesis of proteins and biochemicals necessary for ...
damage, and other
disease A disease is a particular abnormal condition that negatively affects the structure or function of all or part of an organism, and that is not immediately due to any external injury. Diseases are often known to be medical conditions that a ...
s.


See also

*
Auxin binding protein In molecular biology, the auxin binding protein family is a family of proteins which bind auxin. They are located in the lumen of the endoplasmic reticulum (ER). The primary structure of these proteins contains an N-terminal hydrophobic leader ...
* Fusicoccin *
Herbicide Herbicides (, ), also commonly known as weedkillers, are substances used to control undesired plants, also known as weeds.EPA. February 201Pesticides Industry. Sales and Usage 2006 and 2007: Market Estimates. Summary in press releasMain page fo ...
; ''specifically, see the section: Auxin'' **
Phenoxy herbicide Phenoxy herbicides (or "phenoxies") are two families of chemicals that have been developed as commercially important herbicides, widely used in agriculture. They share the part structure of phenoxyacetic acid. Auxins The first group to be discover ...
*
Pruning fruit trees Fruit tree pruning is the pruning, cutting and removing of selected parts of a fruit tree. It spans a number of horticulture, horticultural techniques. Pruning often means cutting branches back, sometimes removing smaller limbs entirely. It may ...
*
Tropism A tropism is a biological phenomenon, indicating growth or turning movement of a biological organism, usually a plant, in response to an environmental stimulus. In tropisms, this response is dependent on the direction of the stimulus (as oppos ...
*
Witch's broom Witch's broom or witches' broom is a deformity in a woody plant, typically a tree, where the natural structure of the plant is changed. A dense mass of shoots grows from a single point, with the resulting structure resembling a broom or a ...
*
Toshio Murashige Toshio Murashige is a professor emeritus of University of California Riverside in plant biology. He is most widely known for his efforts in creating the plant tissue culture medium known as Murashige and Skoog medium Murashige and Skoog medium (o ...
* Folke K. Skoog * Kenneth V. Thimann


References


Further reading

* {{Authority control Plant physiology Auxinic herbicides