Plant hormone
Encyclopedia
Plant hormones are chemicals that regulate plant
growth, which, in the UK, are termed 'plant growth substances'. Plant hormones are signal molecule
s produced within the plant
, and occur in extremely low concentration
s. 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 animal
s, lack gland
s 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.
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 affect plant growth. A large number of related chemical compounds are synthesize
d by humans. They are used to regulate the growth of cultivated plants, weed
s, 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 nutrient
s, 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 produce hormones that affect even different regions of the cell producing the hormone.
Hormones are transported within the plant by utilizing four types of movements. For localized movement, cytoplasm
ic streaming within cells and slow diffusion of ion
s and molecule
s 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 sugar
s from the leaves to the root
s 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 meristem
s, 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 carbohydrate
s, amino acid
s, or peptide
s. 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.
The five major classes are:
also called ABA, was discovered and researched under two different names before its chemical properties were fully known, it was called dormin and abscicin II. Once it was determined that the two latter compounds are the same, it was named abscisic acid. The name "abscisic acid" was given because it was 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 chloroplast
s, especially when plants are under stress. In general, it acts as an inhibitory chemical compound that affects bud
growth, and seed and bud dormancy. It mediates changes within the apical meristem, causing bud dormancy and the alteration of the last set of leaves into protective bud covers. Since it was found in freshly abscissed leaves, it was thought to play a role in the processes of natural leaf drop, but further research has disproven this. In plant species from temperate parts of the world, it plays a role in leaf and seed dormancy by inhibiting growth, but, as it is dissipated from seeds or buds, growth begins. In other plants, as ABA levels decrease, growth then commences as gibberellin
levels increase. Without ABA, buds and seeds would start to grow during warm periods in winter and be killed when it froze again. Since ABA dissipates slowly from the tissues and its effects take time to be offset by other plant hormones, there is a delay in physiological pathways that provide some protection from premature growth. It accumulates within seeds during fruit maturation, preventing seed germination within the fruit, or seed germination before winter. Abscisic acid's effects are degraded within plant tissues during cold temperatures or by its removal by water washing in out of the tissues, releasing the seeds and buds from dormancy.
In plants under water stress, ABA plays a role in closing the stomata. Soon after plants are water-stressed and the roots are deficient in water, a signal moves up to the leaves, causing the formation of ABA precursors there, which then move to the roots. The roots then release ABA, which is translocated to the foliage through the vascular system and modulates the potassium and sodium uptake within the guard cell
s, which then lose turgidity, closing the stomata.
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. 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, 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.
s are compounds that positively influence cell enlargement, bud formation and root initiation. They also promote the production of other hormones and in conjunction with cytokinin
s, they control the growth of stems, roots, and fruits, and convert stems into flowers. Auxins were the first class of growth regulators discovered. They affect cell elongation by altering cell wall plasticity. They stimulate cambium
, a subtype of meristem
cells, to divide and in stems cause secondary xylem to differentiate. Auxins act to inhibit the growth of buds lower down the stems (apical dominance
), and also to promote lateral and adventitious root development and growth. Leaf abscission is initiated by the growing point of a plant ceasing to produce auxins. Auxins in seeds regulate specific protein synthesis, as they develop within the flower after pollination
, causing the flower to develop a fruit to contain the developing seeds. Auxins are toxic to plants in large concentrations; they are most toxic to dicots and less so to monocots. Because of this property, synthetic
auxin herbicides including 2,4-D and 2,4,5-T have been developed and used for weed
control. Auxins, especially 1-Naphthaleneacetic acid
(NAA) and Indole-3-butyric acid
(IBA), are also commonly applied to stimulate root growth when taking cuttings of plants. The most common auxin found in plants is indoleacetic acid or IAA. The correlation of auxins and cytokinin
s in the plants is a constant (A/C = const.).
s or CKs are a group of chemicals that influence cell division and shoot formation. They were called kinins in the past when the first cytokinins were isolated from yeast cells. They also help delay senescence or the aging of tissues, are responsible for mediating auxin transport throughout the plant, and affect internodal length and leaf growth. They have a highly synergistic effect in concert with auxins, 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; they in conjunction with ethylene promote abscission of leaves, flower parts, and fruits.
The correlation of auxins and cytokinins in the plants is a constant (A/C = const.).
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 (see Hyponastic response
). 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 stem's 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.
s, 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.
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.
from cutting
s, 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.
Embryo dormancy is characterized by a high ABA:GA ratio, whereas the seed has a high ABA 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.
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
growth, which, in the UK, are termed 'plant growth substances'. Plant hormones are signal molecule
Molecule
A molecule is an electrically neutral group of at least two atoms held together by covalent chemical bonds. Molecules are distinguished from ions by their electrical charge...
s produced within the plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...
, and occur in extremely low concentration
Concentration
In chemistry, concentration is defined as the abundance of a constituent divided by the total volume of a mixture. Four types can be distinguished: mass concentration, molar concentration, number concentration, and volume concentration...
s. Hormones regulate cellular processes in targeted cells
Cell (biology)
The cell is the basic structural and functional unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often called the building block of life. The Alberts text discusses how the "cellular building blocks" move to shape developing embryos....
locally and, when moved to other locations, in other locations of the plant. Hormones also determine the formation of flowers, stems
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...
, leaves
Leaves
-History:Vocalist Arnar Gudjonsson was formerly the guitarist with Mower, and he was joined by Hallur Hallsson , Arnar Ólafsson , Bjarni Grímsson , and Andri Ásgrímsson . Late in 2001 they played with Emiliana Torrini and drew early praise from the New York Times...
, the shedding of leaves
Leaves
-History:Vocalist Arnar Gudjonsson was formerly the guitarist with Mower, and he was joined by Hallur Hallsson , Arnar Ólafsson , Bjarni Grímsson , and Andri Ásgrímsson . Late in 2001 they played with Emiliana Torrini and drew early praise from the New York Times...
, and the development and ripening of fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...
. Plants, unlike animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
s, lack gland
Gland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...
s 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
Senescence
Senescence or biological aging is the change in the biology of an organism as it ages after its maturity. Such changes range from those affecting its cells and their function to those affecting the whole organism...
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
Longevity
The word "longevity" is sometimes used as a synonym for "life expectancy" in demography or known as "long life", especially when it concerns someone or something lasting longer than expected ....
, and even plant death
Death
Death is the permanent termination of the biological functions that sustain a living organism. Phenomena which commonly bring about death include old age, predation, malnutrition, disease, and accidents or trauma resulting in terminal injury....
. Hormones are vital
Vital
Vital or Vitals may refer to:* VITAL for Children, a charitable organisation* Vitalism, the doctrine that life cannot be explained solely by mechanism* Vitalism , the Jain teacher Mahāvīra's philosophy...
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 expressionGene expression
Gene expression is the process by which information from a gene is used in the synthesis of a functional gene product. These products are often proteins, but in non-protein coding genes such as ribosomal RNA , transfer RNA or small nuclear RNA genes, the product is a functional RNA...
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 affect plant growth. A large number of related chemical compounds are synthesize
Chemical synthesis
In chemistry, chemical synthesis is purposeful execution of chemical reactions to get a product, or several products. This happens by physical and chemical manipulations usually involving one or more reactions...
d by humans. They are used to regulate the growth of cultivated plants, weed
Weed
A weed in a general sense is a plant that is considered by the user of the term to be a nuisance, and normally applied to unwanted plants in human-controlled settings, especially farm fields and gardens, but also lawns, parks, woods, and other areas. More specifically, the term is often used to...
s, and in vitro
In vitro
In vitro refers to studies in experimental biology that are conducted using components of an organism that have been isolated from their usual biological context in order to permit a more detailed or more convenient analysis than can be done with whole organisms. Colloquially, these experiments...
-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 nutrient
Nutrient
A nutrient is a chemical that an organism needs to live and grow or a substance used in an organism's metabolism which must be taken in from its environment. They are used to build and repair tissues, regulate body processes and are converted to and used as energy...
s, but chemicals that in small amounts promote and influence the growth, development, and differentiation of cells and tissues
Tissue (biology)
Tissue is a cellular organizational level intermediate between cells and a complete organism. A tissue is an ensemble of cells, not necessarily identical, but from the same origin, that together carry out a specific function. These are called tissues because of their identical functioning...
. 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
Heart
The heart is a myogenic muscular organ found in all animals with a circulatory system , that is responsible for pumping blood throughout the blood vessels by repeated, rhythmic contractions...
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 produce hormones that affect even different regions of the cell producing the hormone.
Hormones are transported within the plant by utilizing four types of movements. For localized movement, cytoplasm
Cytoplasm
The cytoplasm is a small gel-like substance residing between the cell membrane holding all the cell's internal sub-structures , except for the nucleus. All the contents of the cells of prokaryote organisms are contained within the cytoplasm...
ic streaming within cells and slow diffusion of ion
Ion
An ion is an atom or molecule in which the total number of electrons is not equal to the total number of protons, giving it a net positive or negative electrical charge. The name was given by physicist Michael Faraday for the substances that allow a current to pass between electrodes in a...
s and molecule
Molecule
A molecule is an electrically neutral group of at least two atoms held together by covalent chemical bonds. Molecules are distinguished from ions by their electrical charge...
s 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 sugar
Sugar
Sugar is a class of edible crystalline carbohydrates, mainly sucrose, lactose, and fructose, characterized by a sweet flavor.Sucrose in its refined form primarily comes from sugar cane and sugar beet...
s from the leaves to the root
Root
In vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...
s and flowers, and xylem
Xylem
Xylem is one of the two types of transport tissue in vascular plants. . The word xylem is derived from the Classical Greek word ξυλον , meaning "wood"; the best-known xylem tissue is wood, though it is found throughout the plant...
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 meristem
Meristem
A meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....
s, 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 carbohydrate
Carbohydrate
A carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...
s, amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...
s, or peptide
Peptide
Peptides are short polymers of amino acid monomers linked by peptide bonds. They are distinguished from proteins on the basis of size, typically containing less than 50 monomer units. The shortest peptides are dipeptides, consisting of two amino acids joined by a single peptide bond...
s. 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
Mole (unit)
The mole is a unit of measurement used in chemistry to express amounts of a chemical substance, defined as an amount of a substance that contains as many elementary entities as there are atoms in 12 grams of pure carbon-12 , the isotope of carbon with atomic weight 12. This corresponds to a value...
/L
Litér
- External links :*...
). 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
In vitro
In vitro refers to studies in experimental biology that are conducted using components of an organism that have been isolated from their usual biological context in order to permit a more detailed or more convenient analysis than can be done with whole organisms. Colloquially, these experiments...
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
Abscisic acidAbscisic acid
Abscisic acid , also known as abscisin II and dormin, is a plant hormone. ABA functions in many plant developmental processes, including bud dormancy. It is degraded by the enzyme -abscisic acid 8'-hydroxylase.-Function:...
also called ABA, was discovered and researched under two different names before its chemical properties were fully known, it was called dormin and abscicin II. Once it was determined that the two latter compounds are the same, it was named abscisic acid. The name "abscisic acid" was given because it was 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 chloroplast
Chloroplast
Chloroplasts are organelles found in plant cells and other eukaryotic organisms that conduct photosynthesis. Chloroplasts capture light energy to conserve free energy in the form of ATP and reduce NADP to NADPH through a complex set of processes called photosynthesis.Chloroplasts are green...
s, especially when plants are under stress. In general, it acts as an inhibitory chemical compound that affects bud
Bud
In botany, a bud is an undeveloped or embryonic shoot and normally occurs in the axil of a leaf or at the tip of the stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be specialized to develop flowers or short shoots, or may have...
growth, and seed and bud dormancy. It mediates changes within the apical meristem, causing bud dormancy and the alteration of the last set of leaves into protective bud covers. Since it was found in freshly abscissed leaves, it was thought to play a role in the processes of natural leaf drop, but further research has disproven this. In plant species from temperate parts of the world, it plays a role in leaf and seed dormancy by inhibiting growth, but, as it is dissipated from seeds or buds, growth begins. In other plants, as ABA levels decrease, growth then commences as gibberellin
Gibberellin
Gibberellins are plant hormones that regulate growth and influence various developmental processes, including stem elongation, germination, dormancy, flowering, sex expression, enzyme induction, and leaf and fruit senescence....
levels increase. Without ABA, buds and seeds would start to grow during warm periods in winter and be killed when it froze again. Since ABA dissipates slowly from the tissues and its effects take time to be offset by other plant hormones, there is a delay in physiological pathways that provide some protection from premature growth. It accumulates within seeds during fruit maturation, preventing seed germination within the fruit, or seed germination before winter. Abscisic acid's effects are degraded within plant tissues during cold temperatures or by its removal by water washing in out of the tissues, releasing the seeds and buds from dormancy.
In plants under water stress, ABA plays a role in closing the stomata. Soon after plants are water-stressed and the roots are deficient in water, a signal moves up to the leaves, causing the formation of ABA precursors there, which then move to the roots. The roots then release ABA, which is translocated to the foliage through the vascular system and modulates the potassium and sodium uptake within the guard cell
Guard cell
Guard cells are specialized cells located in the Leaf epidermis of plants. Pairs of guard cells surround tiny stomatal airway pores . These tiny holes in the surface of leaves are necessary for gas exchange into and out of the plant; carbon dioxide enters the plant allowing the carbon fixation...
s, which then lose turgidity, closing the stomata.
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
Catabolism
Catabolism is the set of metabolic pathways that break down molecules into smaller units and release energy. In catabolism, large molecules such as polysaccharides, lipids, nucleic acids and proteins are broken down into smaller units such as monosaccharides, fatty acids, nucleotides, and amino...
, within the plant affects metabolic reactions and cellular growth and production of other hormones. 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, 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.
Auxins
AuxinAuxin
Auxins are a class of plant hormones with some morphogen-like characteristics. Auxins have a cardinal role in coordination of many growth and behavioral processes in the plant's life cycle and are essential for plant body development. Auxins and their role in plant growth were first described by...
s are compounds that positively influence cell enlargement, bud formation and root initiation. They also promote the production of other hormones and in conjunction with cytokinin
Cytokinin
Cytokinins are a class of plant growth substances 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 leaf senescence...
s, they control the growth of stems, roots, and fruits, and convert stems into flowers. Auxins were the first class of growth regulators discovered. They affect cell elongation by altering cell wall plasticity. They stimulate cambium
Cambium (botany)
A cambium , in botany, is a tissue layer that provide undifferentiated cells for plant growth. It forms parallel rows of cells, which result in secondary tissues....
, a subtype of meristem
Meristem
A meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....
cells, to divide and in stems cause secondary xylem to differentiate. Auxins act to inhibit the growth of buds lower down the stems (apical dominance
Apical dominance
In plant physiology, apical dominance is the phenomenon whereby the main central stem of the plant is dominant over other side stems; on a branch the main stem of the branch is further dominant over its own side branchlets....
), and also to promote lateral and adventitious root development and growth. Leaf abscission is initiated by the growing point of a plant ceasing to produce auxins. Auxins in seeds regulate specific protein synthesis, as they develop within the flower after pollination
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...
, causing the flower to develop a fruit to contain the developing seeds. Auxins are toxic to plants in large concentrations; they are most toxic to dicots and less so to monocots. Because of this property, synthetic
Chemical synthesis
In chemistry, chemical synthesis is purposeful execution of chemical reactions to get a product, or several products. This happens by physical and chemical manipulations usually involving one or more reactions...
auxin herbicides including 2,4-D and 2,4,5-T have been developed and used for weed
Weed
A weed in a general sense is a plant that is considered by the user of the term to be a nuisance, and normally applied to unwanted plants in human-controlled settings, especially farm fields and gardens, but also lawns, parks, woods, and other areas. More specifically, the term is often used to...
control. Auxins, especially 1-Naphthaleneacetic acid
1-Naphthaleneacetic acid
1-Naphthaleneacetic acid, commonly abbreviated NAA, is an organic compound with the formula C10H7CH2CO2H. This colourless solid is soluble in organic solvents...
(NAA) and Indole-3-butyric acid
Indole-3-butyric acid
Indole-3-butyric acid 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.-Plant hormone:...
(IBA), are also commonly applied to stimulate root growth when taking cuttings of plants. The most common auxin found in plants is indoleacetic acid or IAA. The correlation of auxins and cytokinin
Cytokinin
Cytokinins are a class of plant growth substances 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 leaf senescence...
s in the plants is a constant (A/C = const.).
Cytokinins
CytokininCytokinin
Cytokinins are a class of plant growth substances 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 leaf senescence...
s or CKs are a group of chemicals that influence cell division and shoot formation. They were called kinins in the past when the first cytokinins were isolated from yeast cells. They also help delay senescence or the aging of tissues, are responsible for mediating auxin transport throughout the plant, and affect internodal length and leaf growth. They have a highly synergistic effect in concert with auxins, 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; they in conjunction with ethylene promote abscission of leaves, flower parts, and fruits.
The correlation of auxins and cytokinins in the plants is a constant (A/C = const.).
Ethylene
EthyleneEthylene
Ethylene is a gaseous organic compound with the formula . It is the simplest alkene . Because it contains a carbon-carbon double bond, ethylene is classified as an unsaturated hydrocarbon. Ethylene is widely used in industry and is also a plant hormone...
is a gas that forms through the Yang Cycle from the breakdown of methionine
Methionine
Methionine is an α-amino acid with the chemical formula HO2CCHCH2CH2SCH3. This essential amino acid is classified as nonpolar. This amino-acid is coded by the codon AUG, also known as the initiation codon, since it indicates mRNA's coding region where translation into protein...
, 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 (see Hyponastic response
Hyponastic response
The hyponastic response is an upward bending of leaves or other plant parts, resulting from growth of the lower side. This can be observed in many terrestrial plants and is thought to be linked to the plant hormone ethylene....
). As the new shoot is exposed to light, reactions by phytochrome
Phytochrome
Phytochrome is a photoreceptor, a pigment that plants use to detect light. It is sensitive to light in the red and far-red region of the visible spectrum. Many flowering plants use it to regulate the time of flowering based on the length of day and night and to set circadian rhythms...
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 stem's 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
Climacteric (biology)
In current practice, climacteric is most often a synonym for female menopause. In Princeton University's online Wordnet database, climacteric is listed as: * 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
GibberellinGibberellin
Gibberellins are plant hormones that regulate growth and influence various developmental processes, including stem elongation, germination, dormancy, flowering, sex expression, enzyme induction, and leaf and fruit senescence....
s, 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
Gibberella fujikuroi
Gibberella fujikuroi is a fungal plant pathogen. It causes bakanae disease in rice seedlings, by overloading them with the phytohormone gibberellin as its own metabolic byproduct.- External links :* * *...
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
Endosperm
Endosperm is the tissue produced inside the seeds of most flowering plants around the time of fertilization. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This makes endosperm an important source of nutrition in human diet...
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.
Other known hormones
Other identified plant growth regulators include:- BrassinosteroidBrassinosteroidBrassinosteroids are a class of polyhydroxysteroids that have been recognized as a sixth class ofplant hormones. These were first explored nearly forty years ago when Mitchell et al. reported promotion in stem elongation and cell division by the treatment of organic extracts of rapeseed pollen...
s, are a class of polyhydroxysteroids, a group of plant growth regulators. Brassinosteroids have been recognized as a sixth class of plant hormones, which stimulate cell elongation and division, gravitropismGravitropismGravitropism is a turning or growth movement by a plant or fungus in response to gravity. Charles Darwin was one of the first to scientifically document that roots show positive gravitropism and stems show negative gravitropism. That is, roots grow in the direction of gravitational pull and stems...
, resistance to stress, and xylemXylemXylem is one of the two types of transport tissue in vascular plants. . The word xylem is derived from the Classical Greek word ξυλον , meaning "wood"; the best-known xylem tissue is wood, though it is found throughout the plant...
differentiation. They inhibit rootRootIn vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...
growth and leaf abscissionAbscissionAbscission is a term used in several areas of biology. In plant sciences it most commonly refers to the process by which a plant drops one or more of its parts, such as a leaf, fruit, flower or seed...
. BrassinolideBrassinolideBrassinolide is a plant hormone. The first isolated brassinosteroid, it was discovered when it was shown that pollen from rapeseed could promote stem elongation and cell division. The biologically active component was isolated and named brassinolide....
was the first identified brassinosteroid and was isolated from organic extracts of rapeseed (Brassica napus) pollen in 1970. - Salicylic acidSalicylic acidSalicylic acid is a monohydroxybenzoic acid, a type of phenolic acid and a beta hydroxy acid. This colorless crystalline organic acid is widely used in organic synthesis and functions as a plant hormone. It is derived from the metabolism of salicin...
— activates genes in some plants that produce chemicals that aid in the defense against pathogenic invaders. - Jasmonates — are produced from fatty acids and seem to promote the production of defense proteins that are used to fend off invading organisms. They are believed to also have a role in seed germination, and affect the storage of protein in seeds, and seem to affect root growth.
- Plant peptide hormonesPlant peptide hormonesPeptide signaling plays a significant role in various aspects of plant growth and development and specific receptors for various peptides have been identified as being membrane-localized receptor kinases, the largest family of receptor-like molecules in plants...
— 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. - PolyaminePolyamineA polyamine is an organic compound having two or more primary amino groups .This class of compounds includes several synthetic substances that are important feedstocks for the chemical industry, such as ethylene diamine , 1,3-diaminopropane , and hexamethylenediamine...
s — 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. - Nitric oxideNitric oxideNitric oxide, also known as nitrogen monoxide, is a diatomic molecule with chemical formula NO. It is a free radical and is an important intermediate in the chemical industry...
(NO) — serves as signal in hormonal and defense responses. - StrigolactoneStrigolactoneStrigolactones are plant hormones that have been implicated in inhibition of shoot branching. Strigolactones are carotenoid-derived and trigger germination of parasitic plant seeds and stimulate symbiotic mycorrhizal fungi...
s, implicated in the inhibition of shoot branching. - KarrikinKarrikinKarrikins are a group of plant growth regulators found in the smoke of burning plant material. For many years scientists have known that smoke from forest fires had the ability to stimulate the germination of seeds. In 2004, after studying the thousands of chemical compounds found in smoke, it...
s, a group of plant growth regulators found in the smoke of burning plant material that have the ability to stimulate the germination of seeds.
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 salicylateSodium salicylate
Sodium salicylate is a sodium salt of salicylic acid. It can be prepared from sodium phenolate and carbon dioxide under higher temperature and pressure...
has been found to suppress proliferation of lymphoblastic leukemia, prostate, breast, and melanoma human cancer cells. Jasmonic acid
Jasmonic acid
Jasmonic acid is derived from the fatty acid linolenic acid. It is a member of the jasmonate class of plant hormones. It is biosynthesized from linolenic acid by the octadecanoid pathway....
, 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 propagationPlant propagation
Plant propagation is the process of creating new plants from a variety of sources: seeds, cuttings, bulbs and other plant parts. Plant propagation can also refer to the artificial or natural dispersal of plants.-Sexual propagation :...
from cutting
Cutting
Cutting is the separation of a physical object, or a portion of a physical object, into two portions, through the application of an acutely directed force. An implement commonly used for cutting is the knife or in medical cases the scalpel...
s, grafting
Grafting
Grafting is a horticultural technique whereby tissues from one plant are inserted into those of another so that the two sets of vascular tissues may join together. This vascular joining is called inosculation...
, micropropagation
Micropropagation
Micropropagation is the practice of rapidly multiplying stock plant material to produce a large number of progeny plants, using modern plant tissue culture methods....
, 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 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 a high ABA 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.
External links
- Another quality guide
- Simple plant hormone table with location of synthesis and effects of application — this is the format used in the descriptions at the ends of the Wikipedia articles on individual plant hormones.
- Hormonal Regulation of Gene Expression and Development — Detailed intro including genetic information.