Ethylene (plant hormone)

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An ethylene signal transduction pathway. Ethylene permeates the cell membrane and binds to a gasoreceptor on the endoplasmic reticulum. The gasoreceptor releases the repressed EIN2. This then activates a signal transduction pathway which activates regulatory genes that eventually trigger an ethylene response. The activated DNA is transcribed into mRNA which is then translated into a functional enzyme that is used for ethylene biosynthesis. Ethylene Signal Transduction.svg
An ethylene signal transduction pathway. Ethylene permeates the cell membrane and binds to a gasoreceptor on the endoplasmic reticulum. The gasoreceptor releases the repressed EIN2. This then activates a signal transduction pathway which activates regulatory genes that eventually trigger an ethylene response. The activated DNA is transcribed into mRNA which is then translated into a functional enzyme that is used for ethylene biosynthesis.

Ethylene (CH
2=CH
2) is an unsaturated hydrocarbon gas (alkene) acting as a naturally occurring plant hormone. [1] It is the simplest alkene gas and is the first gas known to act as hormone. [2] It acts at trace levels throughout the life of the plant by stimulating or regulating the ripening of fruit, the opening of flowers, the abscission (or shedding) of leaves [3] and, in aquatic and semi-aquatic species, promoting the 'escape' from submergence by means of rapid elongation of stems or leaves. [4] This escape response is particularly important in rice farming. [5] Commercial fruit-ripening rooms use "catalytic generators" to make ethylene gas from a liquid supply of ethanol. Typically, a gassing level of 500 to 2,000 ppm is used, for 24 to 48 hours. Care must be taken to control carbon dioxide levels in ripening rooms when gassing, as high temperature ripening (20 °C; 68 °F) [6] has been seen to produce CO2 levels of 10% in 24 hours. [7]

Contents

History

Ethylene has a long history of use in agriculture. Ancient Egyptians would gash figs in order to stimulate ripening (wounding stimulates ethylene production by plant tissues). The ancient Chinese would burn incense in closed rooms to enhance the ripening of pears. In the 19th century, city dwellers noticed that gas leaks from street lights led to stunting of growth, death of flowers and premature leaf fall. [6] In 1874, it was discovered that smoke caused pineapple fields to bloom. Smoke contains ethylene, and once this was realized the smoke was replaced with ethephon or naphthalene acetic acid, which induce ethylene production. [8]

The scientific study of ethylene as a factor in plant physiology started in the late 19th century. In 1896, Russian botanist Dimitry Neljubow studied the response pea to illuminating gas to which they showed movement. He discovered ethylene as the active component in the light source that stimulated pea behaviour. [2] He reported his discovery in 1901. [9] Sarah Doubt also showed in 1917 that ethylene from illuminating gas stimulated abscission. [10] Farmers in Florida would commonly get their crops to ripen in sheds by lighting kerosene lamps, which was originally thought to induce ripening from the heat. In 1924, Frank E. Denny discovered that it was the molecule ethylene emitted by the kerosene lamps that induced the ripening. [11] Reporting in the Botanical Gazette , he wrote:

Ethylene was very effective in bringing about the desired result, concentrations as low as one part (by volume) of ethylene in one million parts of air being sufficient to cause green lemons to turn yellow in about six to ten days... Furthermore, coloring with either ethylene or gas from the kerosene stoves caused the loss of the "buttons" (calyx, receptacle, and a portion of the peduncle)... Yellowing of the ethylene treated fruit became visible about the third or fourth day, and full yellow color was developed in six to ten days. Untreated fruit remained green during the same period of time. [12]

The same year, Denny published the experimental details separately, [13] and also experimentally showed that use of ethylene was more advantageous than that of kerosene. [14] In 1934, British biologist Richard Gane discovered that the chemical constituent in ripe bananas could cause ripening of green bananas, as well as faster growth of pea. He showed that the same growth effect could be induced by ethylene. [15] Reporting in Nature that ripe fruit (in this case Worcester Pearmain apple) produced ethylene he said:

The amount of ethylene produced [by the apple] is very small—perhaps of the order of 1 cubic centimetre during the whole life-history of the fruit; and the cause of its prodigious biological activity in such small concentration is a problem for further research. Its production by apple ceases or is very much reduced in the absence of oxygen. [16]

He subsequently showed that ethylene was produced by other fruits as well, and that obtained from apple could induce seed germination and plant growth in different vegetables (but not in cereals). [17] His conclusions were not universally accepted by other scientists. [2] They became more convincing when William Crocker, Alfred Hitchcock, and Percy Zimmerman reported in 1935 that ethylene acts similar to auxins in causing plant growth and senescence of vegetative tissues. This established that ethylene is a plant hormone. [18] [19]

Ethylene biosynthesis in plants

The Yang cycle Yang-cycle.svg
The Yang cycle

Ethylene is produced from essentially all parts of higher plants, including leaves, stems, roots, flowers, fruits, tubers, and seeds. Ethylene production is regulated by a variety of developmental and environmental factors. During the life of the plant, ethylene production is induced during certain stages of growth such as germination, ripening of fruits, abscission of leaves, and senescence of flowers. Ethylene production can also be induced by a variety of external aspects such as mechanical wounding, environmental stresses, and certain chemicals including auxin and other regulators. [20] The pathway for ethylene biosynthesis is named the Yang cycle after the scientist Shang Fa Yang who made key contributions to elucidating this pathway.

Ethylene is biosynthesized from the amino acid methionine to S-adenosyl-L-methionine (SAM, also called Adomet) by the enzyme Met adenosyltransferase. SAM is then converted to 1-aminocyclopropane-1-carboxylic acid (ACC) by the enzyme ACC synthase (ACS). The activity of ACS determines the rate of ethylene production, therefore regulation of this enzyme is key for the ethylene biosynthesis. The final step requires oxygen and involves the action of the enzyme ACC-oxidase (ACO), formerly known as the ethylene forming enzyme (EFE) [21] . Ethylene biosynthesis can be induced by endogenous or exogenous ethylene. ACC synthesis increases with high levels of auxins, especially indole acetic acid (IAA) and cytokinins.

Ethylene perception in plants

Ethylene is perceived by a family of five transmembrane protein dimers such as the ETR1 protein in Arabidopsis. The genes encoding ethylene gasoreceptors have been cloned in the reference plant Arabidopsis thaliana and many other plants. Ethylene gasoreceptors are encoded by multiple genes in plant genomes. Dominant missense mutations in any of the gene family, which comprises five gasoreceptors in Arabidopsis and at least six in tomato, can confer insensitivity to ethylene. [22] Loss-of-function mutations in multiple members of the ethylene-gasoreceptor family result in a plant that exhibits constitutive ethylene responses. [23] DNA sequences for ethylene gasoreceptors have also been identified in many other plant species and an ethylene binding protein has even been identified in Cyanobacteria. [1]

Ethylene response to salt stress

A large portion of the soil has been affected by over salinity and it has been known to limit the growth of many plants. Globally, the total area of saline soil was 397,000,000 ha and in continents like Africa, it makes up 2 percent of the soil. [24] The amount of soil salinization has reached 19.5% of the irrigated land and 2.1% of the dry-land agriculture around the world. [25] Soil salinization affects the plants using osmotic potential by net solute accumulation. The osmotic pressure in the plant is what maintains water uptake and cell turgor to help with stomatal function and other cellular mechanisms. [25] Over generations, many plant genes have adapted, allowing plants’ phenotypes to change and built distinct mechanisms to counter salinity effects.

The plant hormone ethylene is a combatant for salinity in most plants. Ethylene is known for regulating plant growth and development and adapted to stress conditions through a complex signal transduction pathway. Central membrane proteins in plants, such as ETO2, ERS1 and EIN2, are used for ethylene signaling in many plant growth processes. ETO2, Ethylene overproducer 2, is a protein that, when mutated, will gain a function to continually produce ethylene even when there is no stress condition, causing the plant to grow short and stumpy. ERS1, Ethylene response sensor 1, is activated when ethylene is present in the signaling pathway and when mutated, it loses a function and cannot bind to ethylene. This means a response is never activated and the plant will not be able to cope with the abiotic stress. EIN2, Ethylene insensitive 2, is a protein that activates the pathway and when there is a mutation here the EIN2 will block ethylene stimulation and an ethylene response gene will not be activated. Mutations in these proteins can lead to heightened salt sensitivity and limit plant growth. The effects of salinity have been studied on Arabidopsis plants that have mutated ERS1 and EIN4 proteins. [26] These proteins are used for ethylene signaling again certain stress conditions, such as salt and the ethylene precursor ACC is allowing suppress of any sensitivity to the salt stress. [26] Mutations in these pathways can cause lack of ethylene signaling, causing stunt in plant growth and development.

Environmental and biological triggers of ethylene

Environmental cues such as flooding, drought, chilling, wounding, and pathogen attack can induce ethylene formation in plants. In flooding, roots suffer from lack of oxygen, or anoxia, which leads to the synthesis of 1-aminocyclopropane-1-carboxylic acid (ACC). ACC is transported upwards in the plant and then oxidized in leaves. The ethylene produced causes nastic movements (epinasty) of the leaves, perhaps helping the plant to lose less water in compensation for an increase in resistance to water transport through oxygen-deficient roots. [27]

Corolla senescence

The corolla of a plant refers to its set of petals. Corolla development in plants is broken into phases from anthesis to corolla wilting. The development of the corolla is directed in part by ethylene, though its concentration is highest when the plant is fertilized and no longer requires the production or maintenance of structures and compounds that attract pollinators. [28] [29] The role of ethylene in the developmental cycle is as a hormonal director of senescence in corolla tissue. This is evident as ethylene production and emission are maximized in developmental phases post-pollination, until corolla wilting. [28] Ethylene-directed senescence of corolla tissue can be observed as color change in the corolla or the wilting/ death of corolla tissue. At the chemical level, ethylene mediates the reduction in the amount of fragrance volatiles produced. Fragrance volatiles act mostly by attracting pollinators. Ethylene's role in this developmental scenario is to move the plant away from a state of attracting pollinators, so it also aids in decreasing the production of these volatiles.

Ethylene production in corolla tissue does not directly cause the senescence of corolla tissue, but acts by releasing secondary products that are consistent with tissue ageing. While the mechanism of ethylene-mediated senescence are unclear, its role as a senescence-directing hormone can be confirmed by ethylene-sensitive petunia response to ethylene knockdown. Knockdown of ethylene biosynthesis genes was consistent with increased corolla longevity; inversely, up-regulation of ethylene biosynthesis gene transcription factors were consistent with a more rapid senescence of the corolla. [28]

List of plant responses to ethylene

Commercial issues

Ethylene shortens the shelf life of many fruits by hastening fruit ripening and floral senescence. Ethylene will shorten the shelf life of cut flowers and potted plants by accelerating floral senescence and floral abscission. Flowers and plants which are subjected to stress during shipping, handling, or storage produce ethylene causing a significant reduction in floral display. Flowers affected by ethylene include carnation, geranium, petunia, rose, and many others. [43]

Ethylene can cause significant economic losses for florists, markets, suppliers, and growers. Researchers have developed several ways to inhibit ethylene, including inhibiting ethylene synthesis and inhibiting ethylene perception. Aminoethoxyvinylglycine (AVG), Aminooxyacetic acid (AOA), and silver salts are ethylene inhibitors. [44] [45] Inhibiting ethylene synthesis is less effective for reducing post-harvest losses since ethylene from other sources can still have an effect. By inhibiting ethylene perception, fruits, plants and flowers don't respond to ethylene produced endogenously or from exogenous sources. Inhibitors of ethylene perception include compounds that have a similar shape to ethylene, but do not elicit the ethylene response. One example of an ethylene perception inhibitor is 1-methylcyclopropene (1-MCP).

Commercial growers of bromeliads, including pineapple plants, use ethylene to induce flowering. Plants can be induced to flower either by treatment with the gas in a chamber, or by placing a banana peel next to the plant in an enclosed area.

Chrysanthemum flowering is delayed by ethylene gas, [46] and growers have found that carbon dioxide 'burners' and the exhaust fumes from inefficient glasshouse heaters can raise the ethylene concentration to 0.05 ppmv, causing delay in flowering of commercial crops.

See also

Related Research Articles

<span class="mw-page-title-main">Plant hormone</span> Chemical compounds that regulate plant growth and development

Plant hormones are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, including embryogenesis, the regulation of organ size, pathogen defense, stress tolerance and reproductive development. Unlike in animals each plant cell is capable of producing hormones. Went and Thimann coined the term "phytohormone" and used it in the title of their 1937 book.

<span class="mw-page-title-main">Ripening</span> Process in fruits that causes them to become more palatable

Ripening is a process in fruits that causes them to become more palatable. In general, fruit becomes sweeter, less green, and softer as it ripens. Even though the acidity of fruit increases as it ripens, the higher acidity level does not make the fruit seem tarter. This effect is attributed to the Brix-Acid Ratio. Climacteric fruits ripen after harvesting and so some fruits for market are picked green.

<span class="mw-page-title-main">Auxin</span> Plant hormone

Auxins are a class of plant hormones with some morphogen-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 first described auxins and their role in plant growth in the 1920s. Kenneth V. Thimann became the first to isolate one of these phytohormones and to determine its chemical structure as indole-3-acetic acid (IAA). Went and Thimann co-authored a book on plant hormones, Phytohormones, in 1937.

<span class="mw-page-title-main">Cytokinin</span> Class of plant hormones promoting cell division

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 leaf senescence.

Gibberellins (GAs) are plant hormones that regulate various developmental processes, including stem elongation, germination, dormancy, flowering, flower development, and leaf and fruit senescence. They are one of the longest-known classes of plant hormone. It is thought that the selective breeding of crop strains that were deficient in GA synthesis was one of the key drivers of the "green revolution" in the 1960s, a revolution that is credited to have saved over a billion lives worldwide.

<span class="mw-page-title-main">Jasmonate</span> Lipid-based plant hormones

Jasmonate (JA) and its derivatives are lipid-based plant hormones that regulate a wide range of processes in plants, ranging from growth and photosynthesis to reproductive development. In particular, JAs are critical for plant defense against herbivory and plant responses to poor environmental conditions and other kinds of abiotic and biotic challenges. Some JAs can also be released as volatile organic compounds (VOCs) to permit communication between plants in anticipation of mutual dangers.

<span class="mw-page-title-main">Abscisic acid</span> Plant hormone

Abscisic acid is a plant hormone. ABA functions in many plant developmental processes, including seed and bud dormancy, the control of organ size and stomatal closure. It is especially important for plants in the response to environmental stresses, including drought, soil salinity, cold tolerance, freezing tolerance, heat stress and heavy metal ion tolerance.

<span class="mw-page-title-main">Gravitropism</span> Plant growth in reaction to gravity and bending of leaves and roots

Gravitropism 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 feature of all higher and many lower plants as well as other organisms. 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 grow in the opposite direction. This behavior can be easily demonstrated with any potted plant. When laid onto its side, the growing parts of the stem begin to display negative gravitropism, growing upwards. Herbaceous (non-woody) stems are capable of a degree of actual bending, but most of the redirected movement occurs as a consequence of root or stem growth outside. The mechanism is based on the Cholodny–Went model which was proposed in 1927, and has since been modified. Although the model has been criticized and continues to be refined, it has largely stood the test of time.

<span class="mw-page-title-main">Plant senescence</span> Process of aging in plants

Plant senescence is the process of aging in plants. Plants have both stress-induced and age-related developmental aging. Chlorophyll degradation during leaf senescence reveals the carotenoids, such as anthocyanin and xanthophylls, which are the cause of autumn leaf color in deciduous trees. Leaf senescence has the important function of recycling nutrients, mostly nitrogen, to growing and storage organs of the plant. Unlike animals, plants continually form new organs and older organs undergo a highly regulated senescence program to maximize nutrient export.

<span class="mw-page-title-main">Methyl jasmonate</span> Chemical compound

Methyl jasmonate is a volatile organic compound used in plant defense and many diverse developmental pathways such as seed germination, root growth, flowering, fruit ripening, and senescence. Methyl jasmonate is derived from jasmonic acid and the reaction is catalyzed by S-adenosyl-L-methionine:jasmonic acid carboxyl methyltransferase.

<span class="mw-page-title-main">Abscission</span> Shedding of various parts of an organism

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 tail to evade a predator. In mycology, it is the liberation of a fungal spore. In cell biology, abscission refers to the separation of two daughter cells at the completion of cytokinesis.

Thigmomorphogenesis is the response by plants to mechanical sensation (touch) by altering their growth patterns. In the wild, these patterns can be evinced by wind, raindrops, and rubbing by passing animals.

<span class="mw-page-title-main">Lateral root</span> Plant root

Lateral roots, emerging from the pericycle, extend horizontally from the primary root (radicle) and over time makeup the iconic branching pattern of root systems. They contribute to anchoring the plant securely into the soil, increasing water uptake, and facilitate the extraction of nutrients required for the growth and development of the plant. Lateral roots increase the surface area of a plant's root system and can be found in great abundance in several plant species. In some cases, lateral roots have been found to form symbiotic relationships with rhizobia (bacteria) and mycorrhizae (fungi) found in the soil, to further increase surface area and increase nutrient uptake.

<span class="mw-page-title-main">1-Aminocyclopropane-1-carboxylate synthase</span> Class of enzymes

The enzyme aminocyclopropane-1-carboxylic acid synthase catalyzes the synthesis of 1-Aminocyclopropane-1-carboxylic acid (ACC), a precursor for ethylene, from S-Adenosyl methionine, an intermediate in the Yang cycle and activated methyl cycle and a useful molecule for methyl transfer:

<span class="mw-page-title-main">1-Methylcyclopropene</span> Synthetic plant growth regulator blocking the effects of ethylene (competitive inhibitor)

1-Methylcyclopropene (1-MCP) is a cyclopropene derivative used as a synthetic plant growth regulator. It is structurally related to the natural plant hormone ethylene and it is used commercially to slow down the ripening of fruit and to help maintain the freshness of cut flowers.

Peptide 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. Signaling peptides include members of the following protein families.

Generally, fleshy fruits can be divided into two groups based on the presence or absence of a respiratory increase at the onset of ripening. This respiratory increase—which is preceded, or accompanied, by a rise in ethylene—is called a climacteric, and there are marked differences in the development of climacteric and non-climacteric fruits. Climacteric fruit can be either monocots or dicots and the ripening of these fruits can still be achieved even if the fruit has been harvested at the end of their growth period. Non-climacteric fruits ripen without ethylene and respiration bursts, the ripening process is slower, and for the most part they will not be able to ripen if the fruit is not attached to the parent plant. Examples of climacteric fruits include apples, bananas, melons, apricots, tomatoes, as well as most stone fruits. Non-climacteric fruits on the other hand include citrus fruits, grapes, and strawberries Essentially, a key difference between climacteric and non-climacteric fruits is that climacteric fruits continue to ripen following their harvest, whereas non-climacteric fruits do not. The accumulation of starch over the early stages of climacteric fruit development may be a key issue, as starch can be converted to sugars after harvest.

<span class="mw-page-title-main">Phototropism</span> Growth of a plant in response to a light stimulus

In biology, 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 hormone called auxin that reacts when phototropism occurs. This causes the plant to have elongated cells on the furthest side from the light. Phototropism is one of the many plant tropisms, or movements, which respond to external stimuli. Growth towards a light source is called positive phototropism, while growth away from light is called negative phototropism. Negative phototropism is not to be confused with skototropism, which is defined as the growth towards darkness, whereas negative phototropism can refer to either the growth away from a light source or towards the darkness. Most plant shoots exhibit positive phototropism, and rearrange their chloroplasts in the leaves to maximize photosynthetic energy and promote growth. Some vine shoot tips exhibit negative phototropism, which allows them to grow towards dark, solid objects and climb them. The combination of phototropism and gravitropism allow plants to grow in the correct direction.

Gaseous signaling molecules are gaseous molecules that are either synthesized internally (endogenously) in the organism, tissue or cell or are received by the organism, tissue or cell from outside and that are used to transmit chemical signals which induce certain physiological or biochemical changes in the organism, tissue or cell. The term is applied to, for example, oxygen, carbon dioxide, sulfur dioxide, nitrous oxide, hydrogen cyanide, ammonia, methane, hydrogen, ethylene, etc.

The acid-growth hypothesis is a theory that explains the expansion dynamics of cells and organs in plants. It was originally proposed by Achim Hager and Robert Cleland in 1971. They hypothesized that the naturally occurring plant hormone, auxin (indole-3-acetic acid, IAA), induces H+ proton extrusion into the apoplast. Such derived apoplastic acidification then activates a range of enzymatic reactions which modifies the extensibility of plant cell walls. Since its formulation in 1971, the hypothesis has stimulated much research and debate. Most debates have concerned the signalling role of auxin and the molecular nature of cell wall modification. The current version holds that auxin activates small auxin-up RNA (SAUR) proteins, which in turn regulate protein phosphatases that modulate proton-pump activity. Acid growth is responsible for short-term (seconds to minutes) variation in growth rate, but many other mechanisms influence longer-term growth.

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