Plant hormones (or plant growth regulators, or PGRs) are internally-secreted chemicals in plants that are used for regulating the plants' growth. Plant hormones are signal molecules produced at specific locations, occur in low concentrations, and cause altered processes in target cells at other locations.
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Plant hormones (also known as phytohormones) are chemicals that regulate plant growth, which, in the UK, are termed 'plant growth substances'. Plant hormones are signal molecules produced within the plant, and occur in extremely low concentrations. Hormones regulate cellular processes in targeted cells locally and when moved to other locations, in other locations of the plant. Hormones also determine the formation of flowers, stems, leaves, the shedding of leaves, and the development and ripening of fruit. Plants, unlike animals, lack glands that produce and secrete hormones, instead each cell is capable of producing hormones. Plant hormones shape the plant, affecting seed growth, time of flowering, the sex of flowers, senescence of leaves and fruits. They affect which tissues grow upward and which grow downward, leaf formation and stem growth, fruit development and ripening, plant longevity, and even plant death. Hormones are vital to plant growth and lacking them, plants would be mostly a mass of undifferentiated cells.
Characteristics: The word hormone is derived from Greek, meaning 'set in motion.' Plant hormones affect gene expression and transcription levels, cellular division, and growth. They are naturally produced within plants, though very similar chemicals are produced by fungi and bacteria that can also effect plant growth. A large number of related chemical compounds are synthesized by humans, they are used to regulate the growth of cultivated plants, weeds, and in vitro-grown plants and plant cells; these manmade compounds are called Plant Growth Regulators or PGRs for short. Early in the study of plant hormones, "phytohormone" was the commonly-used term, but its use is less widely applied now.
Plant hormones are not nutrients, but chemicals that in small amounts promote and influence the growth, development, and differentiation of cells and tissues. The biosynthesis of plant hormones within plant tissues is often diffuse and not always localized. Plants lack glands to produce and store hormones, because, unlike animals, which have two circulatory systems (lymphatic and cardiovascular) powered by a heart that moves fluids around the body, plants use more passive means to move chemicals around the plant. Plants utilize simple chemicals as hormones, which move more easily through the plant's tissues. They are often produced and used on a local basis within the plant body, plant cells even produce hormones that affect different regions of the cell producing the hormone.
Hormones are transported within the plant by utilizing four types of movements. For localized movement, cytoplasmic streaming within cells and slow diffusion of ions and molecules between cells are utilized. Vascular tissues are used to move hormones from one part of the plant to another; these include sieve tubes that move sugars from the leaves to the roots and flowers, and xylem that moves water and mineral solutes from the roots to the foliage.
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 meristems, before cells have fully differentiated. After production they are sometimes moved to other parts of the plant where they cause an immediate effect or they can be stored in cells to be released later. Plants use different pathways to regulate internal hormone quantities and moderate their effects; they can regulate the amount of chemicals used to biosynthesize hormones. They can store them in cells, inactivate them, or cannibalise already-formed hormones by conjugating them with carbohydrates, amino acids or peptides. Plants can also break down hormones chemically, effectively destroying them. Plant hormones frequently regulate the concentrations of other plant hormones. Plants also move hormones around the plant diluting their concentrations.
The concentration of hormones required for plant responses are very low (10-6 to 10-5 mol/L). Because of these low concentrations, it has been very difficult to study plant hormones, and only since the late 1970s have scientists been able to start piecing together their effects and relationships to plant physiology. Much of the early work on plant hormones involved studying plants that were genetically deficient in one or involved the use of tissue-cultured plants grown in vitro that were subjected to differing ratios of hormones, and the resultant growth compared. The earliest scientific observation and study dates to the 1880s; the determination and observation of plant hormones and their identification was spread-out over the next 70 years.
Classes of plant hormones: In general, it is accepted that there are five major classes of plant hormones, some of which are made up of many different chemicals that can vary in structure from one plant to the next. The chemicals are each grouped together into one of these classes based on their structural similarities and on their effects on plant physiology. Other plant hormones and growth regulators are not easily grouped into these classes; they exist naturally or are synthesized by humans or other organisms, including chemicals that inhibit plant growth or interrupt the physiological processes within plants. Each class has positive as well as inhibitory functions, and most often work in tandem with each other, with varying ratios of one or more interplaying to affect growth regulation. The five major classes are:
Abscisic acid (ABA), also known as abscisin II and dormin, is a plant hormone. ABA functions in many plant developmental processes, including bud dormancy.
ABA was originally believed to be involved in abscission - this is now known only to be the case in a small number of plants. ABA-mediated signalling also plays an important part in plant responses to environmental stress and plant pathogens. The plant genes for ABA biosynthesis and sequence of the pathway have been elucidated. ABA is also produced by some plant pathogenic fungi via a biosynthetic route different from ABA biosynthesis in plants.
Abscisic acid owes its names to its role in the abscission of plant leaves. In preparation for winter, ABA is produced in terminal buds. This slows plant growth and directs leaf primordia to develop scales to protect the dormant buds during the cold season. ABA also inhibits the division of cells in the vascular cambium, adjusting to cold conditions in the winter by suspending primary and secondary growth.
Abscisic acid is also produced in the roots in response to decreased soil water potential and other situations in which the plant may be under stress. ABA then translocates to the leaves, where it rapidly alters the osmotic potential of stomatal guard cells, causing them to shrink and stomata to close. The ABA-induced stomatal closure reduces transpiration thus preventing further water loss from the leaves in times of low water availability.
Several ABA mutant Arabidopsis thaliana plants have been identified – both those deficient in ABA production and those insensitive to its action. ABA-deficient plants show defects in seed dormancy, germination, stomatal regulation and some mutants show stunted growth and brown/yellow leaves. These mutants reflect the importance of ABA in seed germination and early embryo development.
Auxins are a class of plant growth substance and morphogens (often called phytohormone or plant hormone). Auxins have an essential role in coordination of many growth and behavioral processes in the plant life cycle. Auxins and their role in plant growth were first revealed by the Dutch scientist Frits Went.
Auxins derive their name from the Greek word "auxano" = "I grow/increase". They were the first of the major plant hormones to be discovered.
Their patterns of active transport through the plant are complex. They 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. Thus a plant can (as a whole) react to external conditions and adjust to them, without requiring a nervous system. On the molecular level, auxins have an aromatic ring and a carboxylic acid group.
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.
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.
Cytokinins (CK) are a class of plant growth substances (plant hormones) that promote cell division. They are primarily involved in cell growth, differentiation, and other physiological processes. Their effects were first discovered through the use of coconut milk in the 1940s by a scientist at the University of Wisconsin–Madison named Folke Skoog.
Cytokinins are involved in many plant processes, including cell division, shoot and root morphogenesis, chloroplast maturation, cell enlargement, auxiliary bud release and senescence. The ratio of auxin to cytokinin is crucial during cell division and the differentiation of plant tissues.
While cytokinin action in vascular plants is described as pleiotropic, this class of plant hormones specifically induces the transition from apical growth to growth via a three-faced apical cell in moss protonema. This bud induction can be pinpointed to differentiation of a specific single cell, and thus is a very specific effect of cytokinin.
Ethylene is a gas that forms through the Yang Cycle from the breakdown of methionine, which is in all cells. Ethylene has very limited solubility in water and does not accumulate within the cell but diffuses out of the cell and escapes out of 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. As the new shoot is exposed to light, reactions by phytochrome in the plant's cells produce a signal for ethylene production to decrease, allowing leaf expansion. Ethylene affects cell growth and cell shape; when a growing shoot hits an obstacle while underground, ethylene production greatly increases, preventing cell elongation and causing the stem to swell. The resulting thicker stem can exert more pressure against the object impeding its path to the surface. If the shoot does not reach the surface and the ethylene stimulus becomes prolonged, it affects the stems natural geotropic response, which is to grow upright, allowing it to grow around an object. Studies seem to indicate that ethylene affects stem diameter and height: When stems of trees are subjected to wind, causing lateral stress, greater ethylene production occurs, resulting in thicker, more sturdy tree trunks and branches. Ethylene affects fruit-ripening: Normally, when the seeds are mature, ethylene production increases and builds-up within the fruit, resulting in a climacteric event just before seed dispersal. The nuclear protein Ethylene Insensitive2 (EIN2) is regulated by ethylene production, and, in turn, regulates other hormones including ABA and stress hormones
Gibberellins or 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. Gibberellins are important in seed germination, affecting enzyme production that mobilizes food production used for growth of new cells. This is done by modulating chromosomal transcription. In grain (rice, wheat, corn, etc.) seeds, a layer of cells called the aleurone layer wraps around the endosperm tissue. Absoption of water by the seed causes production of GA. The GA is transported to the aleurone layer, which responds by producing enzymes that break down stored food reserves within the endosperm, which are utilized by the growing seedling. GAs produce bolting of rosette-forming plants, increasing internodal length. They promote flowering, cellular division, and in seeds growth after germination. Gibberellins also reverse the inhibition of shoot growth and dormancy induced by ABA.
Potential medical applications: Plant stress hormones activate cellular responses, including cell death, to diverse stress situations in plants. Researchers have found that some plant stress hormones share the ability to adversely affect human cancer cells . For example, sodium salicylate has been found to suppress proliferation of lymphoblastic leukemia, prostate, breast, and melanoma human cancer cells. Jasmonic acid, a plant stress hormone that belongs to the jasmonate family, induced death in lymphoblastic leukemia cells. Methyl jasmonate has been found to induce cell death in a number of cancer cell lines.
Hormones and plant propagation: Synthetic plant hormones or PGRs are commonly used in a number of different techniques involving plant propagation from cuttings, grafting, micropropagation, and tissue culture. 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.
Seed dormancy: Plant hormones affect seed germinations and dormancy by affecting different parts of the seed. Embryo dormancy is characterized by a high ABA/GA ratio, whereas the seed has a high ABA sensitivity and low GA sensitivity. To release the seed from this type of dormancy and initiate seed germination, an alteration in hormone biosynthesis and degradation towards a low ABA/GA ratio, along with a decrease in ABA sensitivity and an increase in GA sensitivity needs to occur.
Florigen (or flowering hormone) is the term used to describe the hypothesized hormone-like molecules responsible for controlling and/or triggering flowering in plants. Florigen is produced in the leaves and acts in the shoot apical meristem of buds and growing tips. It is known to be graft-transmissible and even functions between species. However despite having been sought since the 1930s, the exact nature of florigen is still a mystery.
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