Plant hormone
Plant hormones (or phytohormones) are
Phytohormones occur across the
Characteristics
The word hormone is derived from Greek, meaning set in motion. Plant hormones affect
Plant hormones are not
Hormones are transported within the plant by utilizing four types of movements. For localized movement,
Not all plant cells respond to hormones, but those cells that do are programmed to respond at specific points in their growth cycle. The greatest effects occur at specific stages during the cell's life, with diminished effects occurring before or after this period. Plants need hormones at very specific times during plant growth and at specific locations. They also need to disengage the effects that hormones have when they are no longer needed. The production of hormones occurs very often at sites of active growth within the
The concentration of hormones required for plant responses are very low (10−6 to 10−5
Synergism in plant hormones refers to the how of two or more hormones result in an effect that is more than the individual effects. For example, auxins and cytokinins often act in cooperation during cellular division and differentiation. Both hormones are key to cell cycle regulation, but when they come together, their synergistic interactions can enhance cell proliferation and organogenesis more effectively than either could in isolation.
Classes
Different hormones can be sorted into different classes, depending on their chemical structures. Within each class of hormone, chemical structures can vary, but all members of the same class have similar physiological effects. Initial research into plant hormones identified five major classes: abscisic acid, auxins, brassinosteroids, cytokinins and ethylene.[16] This list was later expanded, and brassinosteroids, jasmonates, salicylic acid, and strigolactones are now also considered major plant hormones. Additionally there are several other compounds that serve functions similar to the major hormones, but their status as bona fide hormones is still debated.
Abscisic acid
Abscisic acid (also called ABA) is one of the most important plant growth inhibitors. It was discovered and researched under two different names, dormin and abscicin II, before its chemical properties were fully known. Once it was determined that the two compounds are the same, it was named abscisic acid. The name refers to the fact that it is found in high concentrations in newly abscissed or freshly fallen leaves.
This class of PGR is composed of one chemical compound normally produced in the leaves of plants, originating from
ABA exists in all parts of the plant, and its concentration within any tissue seems to mediate its effects and function as a hormone; its degradation, or more properly catabolism, within the plant affects metabolic reactions and cellular growth and production of other hormones.[18] Plants start life as a seed with high ABA levels. Just before the seed germinates, ABA levels decrease; during germination and early growth of the seedling, ABA levels decrease even more. As plants begin to produce shoots with fully functional leaves, ABA levels begin to increase again, slowing down cellular growth in more "mature" areas of the plant. Stress from water or predation affects ABA production and catabolism rates, mediating another cascade of effects that trigger specific responses from targeted cells. Scientists are still piecing together the complex interactions and effects of this and other phytohormones.
In plants under water stress, ABA plays a role in closing the
Auxins
Auxins are compounds that positively influence cell enlargement, bud formation, and root initiation. They also promote the production of other hormones and, in conjunction with cytokinins, control the growth of stems, roots, and fruits, and convert stems into flowers.[22] Auxins were the first class of growth regulators discovered. A Dutch Biologist
Auxins act to inhibit the growth of buds lower down the stems in a phenomenon known as
In large concentrations, auxins are often toxic to plants; they are most toxic to
Brassinosteroids
Cytokinins
Cytokinins or CKs are a group of chemicals that influence cell division and shoot formation. They also help delay senescence of tissues, are responsible for mediating auxin transport throughout the plant, and affect internodal length and leaf growth. They were called kinins in the past when they were first isolated from yeast cells. Cytokinins and auxins often work together, and the ratios of these two groups of plant hormones affect most major growth periods during a plant's lifetime. Cytokinins counter the apical dominance induced by auxins; in conjunction with ethylene, they promote abscission of leaves, flower parts, and fruits.[31]
Among the plant hormones, the three that are known to help with immunological interactions are ethylene (ET), salicylates (SA), and jasmonates (JA), however more research has gone into identifying the role that cytokinins (CK) play in this. Evidence suggests that cytokinins delay the interactions with pathogens, showing signs that they could induce resistance toward these pathogenic bacteria. Accordingly, there are higher CK levels in plants that have increased resistance to pathogens compared to those which are more susceptible.[32] For example, pathogen resistance involving cytokinins was tested using the Arabidopsis species by treating them with naturally occurring CK (trans-zeatin) to see their response to the bacteria Pseudomonas syringa. Tobacco studies reveal that over expression of CK inducing IPT genes yields increased resistance whereas over expression of CK oxidase yields increased susceptibility to pathogen, namely P. syringae.
While there’s not much of a relationship between this hormone and physical plant behavior, there are behavioral changes that go on inside the plant in response to it. Cytokinin defense effects can include the establishment and growth of microbes (delay leaf senescence), reconfiguration of secondary metabolism or even induce the production of new organs such as galls or nodules.[33] These organs and their corresponding processes are all used to protect the plants against biotic/abiotic factors.
Ethylene
Unlike the other major plant hormones, ethylene is a gas and a very simple organic compound, consisting of just six atoms. It forms through the breakdown of methionine, an amino acid which is in all cells. Ethylene has very limited solubility in water and therefore does not accumulate within the cell, typically diffusing out of the cell and escaping the plant. Its effectiveness as a plant hormone is dependent on its rate of production versus its rate of escaping into the atmosphere. Ethylene is produced at a faster rate in rapidly growing and dividing cells, especially in darkness. New growth and newly germinated seedlings produce more ethylene than can escape the plant, which leads to elevated amounts of ethylene, inhibiting leaf expansion (see hyponastic response).
As the new shoot is exposed to light, reactions mediated by
Ethylene also affects fruit ripening. Normally, when the seeds are mature, ethylene production increases and builds up within the fruit, resulting in a
Gibberellins
Gibberellins (GAs) include a large range of chemicals that are produced naturally within plants and by fungi. They were first discovered when Japanese researchers, including Eiichi Kurosawa, noticed a chemical produced by a fungus called Gibberella fujikuroi that produced abnormal growth in rice plants.[40] It was later discovered that GAs are also produced by the plants themselves and control multiple aspects of development across the life cycle. The synthesis of GA is strongly upregulated in seeds at germination and its presence is required for germination to occur. In seedlings and adults, GAs strongly promote cell elongation. GAs also promote the transition between vegetative and reproductive growth and are also required for pollen function during fertilization.[41]
Gibberellins breaks the dormancy (in active stage) in seeds and buds and helps increasing the height of the plant. It helps in the growth of the stem[citation needed]
Jasmonates
JAs have been shown to interact in the signalling pathway of other hormones in a mechanism described as “crosstalk.” The hormone classes can have both negative and positive effects on each other's signal processes.[45]
Jasmonate mutants are more readily consumed by herbivores than wild type plants, indicating that JAs play an important role in the execution of plant defense. When herbivores are moved around leaves of wild type plants, they reach similar masses to herbivores that consume only mutant plants, implying the effects of JAs are localized to sites of herbivory.[48] Studies have shown that there is significant crosstalk between defense pathways.[49]
Salicylic acid
Salicylic acid (SA) is a hormone with a structure related to benzoic acid and phenol. It was originally isolated from an extract of white willow bark (Salix alba) and is of great interest to human medicine, as it is the precursor of the painkiller aspirin. In plants, SA plays a critical role in the defense against biotrophic pathogens. In a similar manner to JA, SA can also become methylated. Like MeJA, methyl salicylate is volatile and can act as a long-distance signal to neighboring plants to warn of pathogen attack. In addition to its role in defense, SA is also involved in the response of plants to abiotic stress, particularly from drought, extreme temperatures, heavy metals, and osmotic stress.[50]
Salicylic acid (SA) serves as a key hormone in plant innate immunity, including resistance in both local and systemic tissue upon biotic attacks, hypersensitive responses, and cell death. Some of the SA influences on plants include seed germination, cell growth, respiration, stomatal closure, senescence-associated gene expression, responses to abiotic and biotic stresses, basal thermo tolerance and fruit yield. A possible role of salicylic acid in signaling disease resistance was first demonstrated by injecting leaves of resistant tobacco with SA.[51] The result was that injecting SA stimulated pathogenesis related (PR) protein accumulation and enhanced resistance to tobacco mosaic virus (TMV) infection. Exposure to pathogens causes a cascade of reactions in the plant cells. SA biosynthesis is increased via isochorismate synthase (ICS) and phenylalanine ammonia-lyase (PAL) pathway in plastids.[52] It was observed that during plant-microbe interactions, as part of the defense mechanisms, SA is initially accumulated at the local infected tissue and then spread all over the plant to induce systemic acquired resistance at non-infected distal parts of the plant. Therefore with increased internal concentration of SA, plants were able to build resistant barriers for pathogens and other adverse environmental conditions[53]
Strigolactones
Strigolactones (SLs) were originally discovered through studies of the germination of the parasitic weed Striga lutea. It was found that the germination of Striga species was stimulated by the presence of a compound exuded by the roots of its host plant.[54] It was later shown that SLs that are exuded into the soil also promote the growth of symbiotic arbuscular mycorrhizal (AM) fungi.[55] More recently, another role of SLs was identified in the inhibition of shoot branching.[56] This discovery of the role of SLs in shoot branching led to a dramatic increase in the interest in these hormones, and it has since been shown that SLs play important roles in leaf senescence, phosphate starvation response, salt tolerance, and light signalling.[57]
Other known hormones
Other identified plant growth regulators include:
- Plant peptide hormones – encompasses all small secreted peptides that are involved in cell-to-cell signaling. These small peptide hormones play crucial roles in plant growth and development, including defense mechanisms, the control of cell division and expansion, and pollen self-incompatibility.[58] The small peptide CLE25 is known to act as a long-distance signal to communicate water stress sensed in the roots to the stomata in the leaves.[59]
- Polyamines – are strongly basic molecules with low molecular weight that have been found in all organisms studied thus far. They are essential for plant growth and development and affect the process of mitosis and meiosis. In plants, polyamines have been linked to the control of senescence[60] and programmed cell death.[61]
- Nitric oxide (NO) – serves as signal in hormonal and defense responses (e.g. stomatal closure, root development, germination, nitrogen fixation, cell death, stress response).[62] NO can be produced by a yet undefined NO synthase, a special type of nitrite reductase, nitrate reductase, mitochondrial cytochrome c oxidase or non enzymatic processes and regulate plant cell organelle functions (e.g. ATP synthesis in chloroplasts and mitochondria).[63]
- Karrikins – are not plant hormones as they are not produced by plants themselves but are rather found in the smoke of burning plant material. Karrikins can promote seed germination in many species.[64] The finding that plants which lack the receptor of karrikin receptor show several developmental phenotypes (enhanced biomass accumulation and increased sensitivity to drought) have led some to speculate on the existence of an as yet unidentified karrikin-like endogenous hormone in plants. The cellular karrikin signalling pathway shares many components with the strigolactone signalling pathway.[65]
- Triacontanol – a fatty alcohol that acts as a growth stimulant, especially initiating new basal breaks in the rose family. It is found in alfalfa(lucerne), bee's wax, and some waxy leaf cuticles.
Use in horticulture
Synthetic plant hormones or PGRs are used in a number of different techniques involving plant propagation from cuttings, grafting, micropropagation and tissue culture. Most commonly they are commercially available as "rooting hormone powder".
The propagation of plants by cuttings of fully developed leaves, stems, or roots is performed by gardeners utilizing auxin as a rooting compound applied to the cut surface; the auxins are taken into the plant and promote root initiation. In grafting, auxin promotes callus tissue formation, which joins the surfaces of the graft together. In micropropagation, different PGRs are used to promote multiplication and then rooting of new plantlets. In the tissue-culturing of plant cells, PGRs are used to produce callus growth, multiplication, and rooting.
When used in field conditions, plant hormones or mixtures that include them can be applied as biostimulants.[66]
Seed dormancy
Plant hormones affect seed germination and dormancy by acting on different parts of the seed.
Embryo dormancy is characterized by a high ABA:GA ratio, whereas the seed has high abscisic acid sensitivity and low GA sensitivity. In order to release the seed from this type of dormancy and initiate seed germination, an alteration in hormone biosynthesis and degradation toward a low ABA/GA ratio, along with a decrease in ABA sensitivity and an increase in GA sensitivity, must occur.
ABA controls embryo dormancy, and GA embryo germination. Seed coat dormancy involves the mechanical restriction of the seed coat. This, along with a low embryo growth potential, effectively produces seed dormancy. GA releases this dormancy by increasing the embryo growth potential, and/or weakening the seed coat so the radical of the seedling can break through the seed coat. Different types of seed coats can be made up of living or dead cells, and both types can be influenced by hormones; those composed of living cells are acted upon after seed formation, whereas the seed coats composed of dead cells can be influenced by hormones during the formation of the seed coat. ABA affects testa or seed coat growth characteristics, including thickness, and effects the GA-mediated embryo growth potential. These conditions and effects occur during the formation of the seed, often in response to environmental conditions. Hormones also mediate endosperm dormancy: Endosperm in most seeds is composed of living tissue that can actively respond to hormones generated by the embryo. The endosperm often acts as a barrier to seed germination, playing a part in seed coat dormancy or in the germination process. Living cells respond to and also affect the ABA:GA ratio, and mediate cellular sensitivity; GA thus increases the embryo growth potential and can promote endosperm weakening. GA also affects both ABA-independent and ABA-inhibiting processes within the endosperm.[67]
Human use
Salicylic acid
Willow bark has been used for centuries as a painkiller. The active ingredient in willow bark that provides these effects is the hormone salicylic acid (SA). In 1899, the pharmaceutical company Bayer began marketing a derivative of SA as the drug aspirin.[68] In addition to its use as a painkiller, SA is also used in topical treatments of several skin conditions, including acne, warts and psoriasis.[69] Another derivative of SA, sodium salicylate has been found to suppress proliferation of lymphoblastic leukemia, prostate, breast, and melanoma human cancer cells.[70]
Jasmonic acid
Jasmonic acid (JA) can induce death in lymphoblastic leukemia cells. Methyl jasmonate (a derivative of JA, also found in plants) has been shown to inhibit proliferation in a number of cancer cell lines,[70] although there is still debate over its use as an anti-cancer drug, due to its potential negative effects on healthy cells.[71]
See also
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External links
- Simple plant hormone table with location of synthesis and effects of application — this is the format used in the description templates at bottom of Wikipedia articles about plant hormones.
- Hormonal Regulation of Gene Expression and Development — Detailed introduction to plant hormones, including genetic information.