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Plant hormone
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Lack of the plant hormone auxin can cause abnormal growth (right)
Plant hormones (also known as phytohormones) are chemicals that regulate plant growth. In the United Kingdom, these 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, moved to other locations, in other functional parts 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. So they are also known as growth factors or growth hormones. The term 'Phytohormone' was coined by Went and Thimann and used in the title of their book in 1937.[1]
Phytohormones are found not only in higher plants but in algae, showing similar functions,[2] and in microorganisms, such as unicellular fungi and bacteria, but in these cases they play no hormonal or other immediate physiological role in the producing organism and can, thus, be regarded as secondary metabolites.[3]
Contents
[hide]
1 Characteristics
2 Classes
2.1 Abscisic acid
2.2 Auxins
2.3 Cytokinins
2.4 Ethylene
2.5 Gibberellins
2.6 Other known hormones
3 Potential medical applications
4 Hormones and plant propagation
4.1 Seed dormancy
5 History
6 References
7 External links
Characteristics[edit]
Phyllody on a purple coneflower (Echinacea purpurea), a plant development abnormality where leaf-like structures replace flower organs. It can be caused by hormonal imbalance, among other reasons.
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 affect plant growth.[4] 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,[5] 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 their bodies. Plants utilize simple chemicals as hormones, which move more easily through their 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, 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 or phloem 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.[6] 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.[7] 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[edit]
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.[8]
The five major classes are:
Abscisic acid[edit]
Abscisic acid (also called ABA) is one of the most important plant growth regulators. It 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 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 chloroplasts, 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.[9]
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[10] and modulates the potassium and sodium uptake within the guard cells, which then lose turgidity, closing the stomata.[11][12] 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.[13] 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[edit]
Main article: Auxin
The auxin indole-3-acetic acid
Auxins are compounds that positively influence cell enlargement, bud formation and root initiation. They also promote the production of other hormones and in conjunction with cytokinins, they control the growth of stems, roots, and fruits, and convert stems into flowers.[14] Auxins were the first class of growth regulators discovered.[15] 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,[16] 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(2,4-dichlorophenoxyacetic) 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 indole-3-acetic acid or IAA. The correlation of auxins and cytokinins in the plants is a constant (A/C = const.)[citation needed].
Cytokinins[edit]
The cytokinin zeatin, the name is derived from Zea maize, in which it was first discovered in immature kernels.
Cytokinins 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 of tissues, are responsible for mediating auxin transport throughout the plant, and affect internodal length and leaf growth. Cytokinins and auxins often work together, and the ratios of these two groups of plant hormones affect most major growth periods during a plant's lifetime. Cytokinins counter the apical dominance induced by auxins; they in conjunction with ethylene promote abscission of leaves, flower parts, and fruits.[17] The correlation of auxins and cytokinins in the plants is a constant (A/C = const.).[citation needed]
Ethylene[edit]
Ethylene
Ethylene is a gas that forms through 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.[18]
Gibberellins[edit]
Main article: Gibberellins
Gibberellin A1
Main function: initiate mobilization of storage materials in seeds during germination, cause elongation of stems, stimulate bolting in biennials, stimulate pollen tube growth.
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.[19] 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. Absorption 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.[20]
Other known hormones[edit]
Other identified plant growth regulators include:
Brassinosteroids – 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, gravitropism, resistance to stress, and xylem differentiation. They inhibit root growth and leaf abscission. Brassinolide was the first identified brassinosteroid and was isolated from extracts of rapeseed (Brassica napus) pollen in 1979.[21]
Salicylic acid – 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 hormones – encompasses all small secreted peptides that are involved in cell-to-cell signaling. These small peptide hormones play crucial roles in plant growth and development, including defense mechanisms, the control of cell division and expansion, and pollen self-incompatibility.[22]
Polyamines – are strongly basic molecules with low molecular weight that have been found in all organisms studied thus far. They are essential for plant growth and development and affect the process of mitosis and meiosis.
Nitric oxide (NO) – serves as signal in hormonal and defense responses (e.g. stomatal closure, root development, germination, nitrogen fixation, cell death, stress response).[23] NO can be produced by a yet undefined NO synthase, a special type of nitrite reductase, nitrate reductase, mitochondrial cytochrome c oxidase or non enzymatic processes and regulate plant cell organelle functions (e.g. ATP synthesis in chloroplasts and mitochondria).[24]
Strigolactones – implicated in the inhibition of shoot branching.[25]
Karrikins – not plant hormones because they are not made by plants, but are a group of plant growth regulators found in the smoke of burning plant material that have the ability to stimulate the germination of seeds [26]
Triacontanol – a fatty alcohol that acts as a growth stimulant, especially initiating new basal breaks in the rose family. It is found in alfalfa (lucerne), bee's wax, and some waxy leaf cuticles.
Potential medical applications[edit]
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.[27] 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[edit]
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[edit]
Main article: 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.[28]
History[edit]
This section needs expansion. You can help by adding to it. (April 2018)
References[edit]
Jump up ^ Went, F.W.; Thimann, K.V. (1937). Phytohormones. New York: The Macmillan Company.
Jump up ^ Tarakhovskaya, E. R.; Maslov, Yu; Shishova, M. F. (2007). "Phytohormones in algae". Russian Journal of Plant Physiology. 54 (2): 163–170. doi:10.1134/s1021443707020021.
Jump up ^ "Gibberellin formation in microorganisms". Plant Growth Regulation. 15: 303–314. doi:10.1007/BF00029903.
Jump up ^ Srivastava, L. M. (2002). Plant growth and development: hormones and environment. Academic Press. p. 140. ISBN 0-12-660570-X.
Jump up ^ Öpik, Helgi; Rolfe, Stephen A.; Willis, Arthur John; Street, Herbert Edward (2005). The physiology of flowering plants (4th ed.). Cambridge University Press. p. 191. ISBN 978-0-521-66251-2.
Jump up ^ Swarup R, Perry P, Hagenbeek D, et al. (July 2007). "Ethylene upregulates auxin biosynthesis in Arabidopsis seedlings to enhance inhibition of root cell elongation". Plant Cell. 19 (7): 2186–96. doi:10.1105/tpc.107.052100. PMC 1955695  . PMID 17630275.
Jump up ^ Srivastava 2002, p. 143
Jump up ^ Weier, Thomas Elliot; Rost, Thomas L.; Weier, T. Elliot (1979). Botany: a brief introduction to plant biology. New York: Wiley. pp. 155–170. ISBN 0-471-02114-8.
Jump up ^ Feurtado JA, Ambrose SJ, Cutler AJ, Ross AR, Abrams SR, Kermode AR (February 2004). "Dormancy termination of western white pine (Pinus monticola Dougl. Ex D. Don) seeds is associated with changes in abscisic acid metabolism". Planta. 218 (4): 630–9. doi:10.1007/s00425-003-1139-8. PMID 14663585.
Jump up ^ Ren H, Gao Z, Chen L, et al. (2007). "Dynamic analysis of ABA accumulation in relation to the rate of ABA catabolism in maize tissues under water deficit". J. Exp. Bot. 58 (2): 211–9. doi:10.1093/jxb/erl117. PMID 16982652. Archived from the original on 2012-07-10.
Jump up ^ Else M.A.; Coupland D.; Dutton L.; Jackson M.B. (January 2001). "Decreased root hydraulic conductivity reduces leaf water potential, initiates stomatal closure, and slows leaf expansion in flooded plants of castor oil (Ricinus communis) despite diminished delivery of ABA from the roots to shoots in xylem sap". Physiologia Plantarum. 111 (1): 46–54. doi:10.1034/j.1399-3054.2001.1110107.x.
Jump up ^ Yan J, Tsuichihara N, Etoh T, Iwai S (October 2007). "Reactive oxygen species and nitric oxide are involved in ABA inhibition of stomatal opening". Plant Cell Environ. 30 (10): 1320–5. doi:10.1111/j.1365-3040.2007.01711.x. PMID 17727421.
Jump up ^ Kermode AR (December 2005). "Role of Abscisic Acid in Seed Dormancy". J Plant Growth Regul. 24 (4): 319–344. doi:10.1007/s00344-005-0110-2.
Jump up ^ Osborne, Daphné J.; McManus, Michael T. (2005). Hormones, signals and target cells in plant development. Cambridge University Press. p. 158. ISBN 978-0-521-33076-3.
Jump up ^ Classification of auxin related compounds based on similarity of their interaction fields: Extension to a new set of compounds Tomic, S.1,2, Gabdoulline, R.R.1, Kojic-Prodic, B.2 and Wade, R.C.11 European Molecular Biology Laboratory, 69012 Heidelberg, Germany 2Institute Rudjer Boskovic, HR-10000 Zagreb, Croatia
Jump up ^ Walz A, Park S, Slovin JP, Ludwig-Müller J, Momonoki YS, Cohen JD (February 2002). "A gene encoding a protein modified by the phytohormone indoleacetic acid". Proc. Natl. Acad. Sci. U.S.A. 99 (3): 1718–23. Bibcode:2002PNAS...99.1718W. doi:10.1073/pnas.032450399. PMC 122257  . PMID 11830675.
Jump up ^ Sipes DL, Einset JW (August 1983). "Cytokinin stimulation of abscission in lemon pistil explants". J Plant Growth Regul. 2 (1–3): 73–80. doi:10.1007/BF02042235.
Jump up ^ Wang Y, Liu C, Li K, et al. (August 2007). "Arabidopsis EIN2 modulates stress response through abscisic acid response pathway". Plant Mol. Biol. 64 (6): 633–44. doi:10.1007/s11103-007-9182-7. PMID 17533512.
Jump up ^ Grennan, Aleel K (2006). "Gibberellin Metabolism Enzymes in Rice". Plant Physiology. 141 (2): 524–6. doi:10.1104/pp.104.900192. PMC 1475483  . PMID 16760495.
Jump up ^ Tsai F-Y.; Lin C.C.; Kao C.H. (January 1997). "A comparative study of the effects of abscisic acid and methyl jasmonate on seedling growth of rice". Plant Growth Regulation. 21 (1): 37–42. doi:10.1023/A:1005761804191.
Jump up ^ Grove, M. D.; Spencer, G. F.; Rohwedder, W. K.; Mandava, N.; Worley, J. F.; Warthen, J. D.; Steffens, G. L.; Flippen-Anderson, J. L.; Cook, J. C. (1979). "Brassinolide, a plant growth-promoting steroid isolated from Brassica napus pollen". Nature. 281 (5728): 216–217. Bibcode:1979Natur.281..216G. doi:10.1038/281216a0.
Jump up ^ Lindsey, Keith; Casson, Stuart; Chilley, Paul. (2002). "Peptides:new signalling molecules in plants". Trends in Plant Science. 7 (2): 78–83. doi:10.1016/S0960-9822(01)00435-3. PMID 11832279.
Jump up ^ Shapiro AD (2005) Nitric oxide signaling in plants. Vitam Horm. 2005;72:339-98.
Jump up ^ Roszer T (2012) Nitric Oxide Synthesis in the Chloroplast. in: Roszer T. The Biology of Subcellular Nitric Oxide. Springer New York, London, Heidelberg. ISBN 978-94-007-2818-9
Jump up ^ Gomez-Roldan V, Fermas S, Brewer PB, et al. (September 2008). "Strigolactone inhibition of shoot branching". Nature. 455 (7210): 189–94. Bibcode:2008Natur.455..189G. doi:10.1038/nature07271. PMID 18690209.
Jump up ^ Chiwocha, Sheila D. S.; Dixon, Kingsley W.; Flematti, Gavin R.; Ghisalberti, Emilio L.; Merritt, David J.; Nelson, David C.; Riseborough, Julie-Anne M.; Smith, Steven M.; Stevens, Jason C. (2009-10-01). "Karrikins: A new family of plant growth regulators in smoke". Plant Science. 177 (4): 252–256. doi:10.1016/j.plantsci.2009.06.007.
Jump up ^ Fingrut O, Flescher E (April 2002). "Plant stress hormones suppress the proliferation and induce apoptosis in human cancer cells". Leukemia. 16 (4): 608–16. doi:10.1038/sj.leu.2402419. PMID 11960340.
Jump up ^ The Seed Biology Place — Seed Dormancy
External links[edit]
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.
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United States presidential line of succession
From Wikipedia, the free encyclopedia
Jump to: navigation, search
The United States presidential line of succession is the order in which persons may become or act as President of the United States if the incumbent President becomes incapacitated, dies, resigns, or is removed from office (by impeachment by the House of Representatives and subsequent conviction by the Senate). The line of succession is set by the United States Constitution and the Presidential Succession Act of 1947[1] as subsequently amended to include newly created cabinet offices. The succession follows the order of Vice President, Speaker of the House of Representatives, President pro tempore of the Senate, and then the heads of federal executive departments who form the Cabinet of the United States. The Cabinet currently has fifteen members, beginning with the Secretary of State, and followed by the rest in the order of their positions' creation. Those heads of department who are ineligible to act as President are also ineligible to succeed the President by succession, for example most commonly if they are not a natural-born U.S. citizen.
Several constitutional law experts have raised questions as to the constitutionality of the provisions that the Speaker of the House and the President pro tempore of the Senate succeed to the presidency,[2] and in 2003 the Continuity of Government Commission raised a number of other issues with the current line of succession.[3]
Contents
[hide]
1 Current order
2 Eligibility
2.1 Eligibility requirements
2.2 Acting officers
3 Motivation for changes to the succession in 1945
4 Constitutional provisions
5 Acting President and President
6 History of succession law set by Congress
6.1 Presidential Succession Act 1792
6.2 Presidential Succession Act 1886
6.3 Presidential Succession Act 1947
6.4 Postmaster General removed 1971
6.5 Secretary of Homeland Security added 2006
7 Successions beyond Vice President
8 Constitutional concerns
9 See also
10 References
11 Further reading
12 External links
Current order[edit]
The current presidential order of succession is as follows:[4]
Key
Eligible Democrat (D)
Eligible Republican (R)
Eligible Independent (I)
Eligible unknown
Not eligible
No.
Office
Current officer
1
Vice President Mike Pence (R)
2
Speaker of the House of Representatives Paul Ryan (R)
3
President pro tempore of the Senate Orrin Hatch (R)
4
Secretary of State Rex Tillerson (R)
5
Secretary of the Treasury Steven Mnuchin (R)
6
Secretary of Defense James Mattis (I)
7
Attorney General Jeff Sessions (R)
8
Secretary of the Interior Ryan Zinke (R)
9
Secretary of Agriculture Sonny Perdue (R)
10
Secretary of Commerce Wilbur Ross (R)
11
Secretary of Labor Alex Acosta (R)
12
Secretary of Health and Human Services Eric Hargan (R)[a]
Acting
13
Secretary of Housing and Urban Development Ben Carson (R)
–
Secretary of Transportation Elaine Chao (R)[b]
14
Secretary of Energy Rick Perry (R)
15
Secretary of Education Betsy DeVos (R)
16
Secretary of Veterans Affairs David Shulkin (I)
17
Secretary of Homeland Security Elaine Duke (I)[a]
Acting
^ Jump up to: a b This listing assumes that acting officers whose prior appointment required Senate confirmation are eligible for the line of succession. See Acting officers below.
Jump up ^ Not a natural-born citizen (acquired U.S. citizenship by naturalization) and thus ineligible for the Presidency.
Cabinet officers are in line according to the chronological order of their department's creation or the department of which their department is the successor (the Department of Defense being successor to the Department of War, and the Department of Health and Human Services being successor to the Department of Health, Education and Welfare).[a]
Eligibility[edit]
Eligibility requirements[edit]
To be eligible to serve as President, a person must be a natural-born U.S. citizen, at least 35 years of age, and a resident within the United States for at least 14 years. These eligibility requirements are specified both in the U.S. Constitution, Article II, Section 1, Clause 5, and in the Presidential Succession Act of 1947 (3 U.S.C. § 19(e)).
Acting officers[edit]
Acting officers may be eligible. In 2009, the Continuity of Government Commission, a private non-partisan think tank, reported,
The language in the current Presidential Succession Act is less clear than that of the 1886 Act with respect to Senate confirmation. The 1886 Act refers to "such officers as shall have been appointed by the advice and consent of the Senate to the office therein named..." The current act merely refers to "officers appointed, by and with the advice and consent of the Senate." Read literally, this means that the current act allows for acting secretaries to be in the line of succession as long as they are confirmed by the Senate for a post (even for example, the second or third in command within a department). It is common for a second in command to become acting secretary when the secretary leaves office. Though there is some dispute over this provision, the language clearly permits acting secretaries to be placed in the line of succession. (We have spoken to acting secretaries who told us they had been placed in the line of succession.)[5]
Motivation for changes to the succession in 1945[edit]
Two months after succeeding Franklin D. Roosevelt, President Harry S. Truman proposed that the Speaker of the House and the President pro tempore of the Senate be granted priority in the line of succession over the Cabinet so as to ensure the President would not be able to appoint his successor to the Presidency.[6]
The Secretary of State and the other Cabinet officials are appointed by the President, while the Speaker of the House and the President pro tempore of the Senate are elected officials. The Speaker is chosen by the U.S. House of Representatives, and every Speaker has been a member of that body for the duration of their term as Speaker, though this is not technically a requirement; the President pro tempore is chosen by the U.S. Senate and customarily the Senator of the majority party with the longest record of continuous service fills this position. The Congress approved this change and inserted the Speaker and President pro tempore in line, ahead of the members of the Cabinet in the order in which their positions were established.
In his speech supporting the changes, Truman noted that the House of Representatives is more likely to be in political agreement with the President and Vice President than the Senate. The succession of a Republican to a Democratic Presidency would further complicate an already unstable political situation. However, when the changes to the succession were signed into law, they placed Republican House Speaker Joseph W. Martin first in the line of succession after the Vice President.[7]
Some of Truman's critics said that his ordering of the Speaker before the President pro tempore was motivated by his dislike of the then-current holder of the latter rank, Senator Kenneth McKellar.[8] Further motivation may have been provided by Truman's preference for House Speaker Sam Rayburn to be next in the line of succession, rather than Secretary of State Edward R. Stettinius, Jr.[7]
Constitutional provisions[edit]
The line of succession is mentioned in three places in the Constitution:
Article II, Section 1, Clause 6 makes the Vice President first in the line of succession and allows the Congress to provide by law for cases in which neither the President nor Vice President can serve. The current such law governing succession is the Presidential Succession Act of 1947 (3 U.S.C. § 19).
Section 3 of the 20th Amendment provides that if the President-elect dies before his term begins, the Vice President-elect becomes President on Inauguration Day and serves for the full term to which the President-elect was elected. The section also provides that if, on Inauguration Day, a president has not been chosen or the President-elect does not qualify for the presidency, the Vice President-elect acts as president until a president is chosen or the President-elect qualifies. Finally, Section 3 allows the Congress to provide by law for cases in which neither a President-elect nor a Vice President-elect is eligible or available to serve.
The 25th Amendment, ratified in 1967, clarified Article II, Section 1: that the Vice President is the direct successor of the President. He or she becomes President if the President dies, resigns or is removed from office. The amendment also provides for the situation where the President is temporarily disabled, such as if the President has a surgical procedure or becomes mentally unfit.
The amendment also provides for Vice Presidential succession, by requiring Vice Presidential vacancies to be filled by the President and confirmed by both houses of the Congress. Previously, whenever a Vice President had succeeded to the Presidency or had otherwise left the office empty (through death, resignation, or removal from office), the Vice Presidency remained vacant until the next Presidential and Vice Presidential terms began.
Acting President and President[edit]
Main article: Acting President of the United States
New President Lyndon Johnson was sworn in aboard Air Force One, following the assassination of John F. Kennedy in 1963.
Article II, Section 1 of the United States Constitution provides that:
In case of the removal of the President from office, or of his death, resignation, or inability to discharge the powers and duties of the said office, the same shall devolve on the Vice President ... until the disability be removed, or a President elected.
This originally left open the question whether "the same" refers to "the said office" or only "the powers and duties of the said office." Some historians, including Edward Corwin[9] and John D. Feerick,[9] have argued that the framers' intention was that the Vice President would remain Vice President while executing the powers and duties of the Presidency; however, there is also much evidence to the contrary, the most compelling of which is Article I, section 3, of the Constitution itself, the relevant text of which reads:
The Vice President of the United States shall be President of the Senate, but shall have no vote, unless they be equally divided.
The Senate shall chuse [sic] their other officers, and also a President pro tempore, in the absence of the Vice President, or when he shall exercise the office of President of the United States.
This text appears to answer the hypothetical question of whether the office or merely the powers of the Presidency devolved upon the Vice President on his succession. Thus, the 25th Amendment merely restates and reaffirms the validity of existing precedent, apart from adding new protocols for Presidential disability.[original research?] Not everyone agreed with this interpretation when it was first actually tested, and it was left to Vice President John Tyler, the first presidential successor in U.S. history, to establish the precedent that was respected in the absence of the 25th Amendment.
In 1841, John Tyler became the first person to succeed to the presidency.
Upon the death of President William Henry Harrison in 1841 and after a brief hesitation, Tyler took the position that he was the President and not merely acting President upon taking the presidential oath of office. However, some contempories—including John Quincy Adams,[9][10] Henry Clay[11] and other members of Congress[10][11] Whig party leaders[11] and even Tyler's own cabinet[10][11]—believed that he was only acting as President and did not have the office itself.
Nonetheless, Tyler adhered to his position, even returning, unopened, mail addressed to the "Acting President of the United States" sent by his detractors.[12] Tyler's view ultimately prevailed when the Senate voted to accept the title "President,"[11] and this precedent was followed thereafter. The question was finally resolved by Section 1 of the 25th Amendment, which specifies that "In case of the removal of the President from office or of his death or resignation, the Vice President shall become President." The Amendment does not specify whether officers other than the Vice President can become President rather than Acting President in the same set of circumstances. The Presidential Succession Act refers only to other officers acting as President rather than becoming President.
History of succession law set by Congress[edit]
Presidential Succession Act 1792[edit]
Main article: Presidential Succession Act
Wikisource has original text related to this article:
Presidential Succession Act 1792
The Presidential Succession Act of 1792 was the first succession law passed by Congress. The act was contentious because the Federalists did not want the then Secretary of State, Thomas Jefferson, who had become the leader of the Democratic-Republicans, to follow the Vice President in the succession. There were also separation of powers concerns over including the Chief Justice of the United States in the line. The compromise they worked out established the President pro tempore of the Senate as next in line after the Vice President, followed by the Speaker of the House of Representatives.
In either case, these officers were to "act as President of the United States until the disability be removed or a president be elected." The Act called for a special election to be held in November of the year in which dual vacancies occurred (unless the vacancies occurred after the first Wednesday in October, in which case the election would occur the following year; or unless the vacancies occurred within the last year of the presidential term, in which case the next election would take place as regularly scheduled). The people elected President and Vice President in such a special election would have served a full four-year term beginning on March 4 of the next year, but no such election ever took place.
Presidential Succession Act 1886[edit]
Wikisource has original text related to this article:
Presidential Succession Act 1886
In 1881, after the death of President Garfield, and in 1885, after the death of Vice President Hendricks, there had been no President pro tempore in office, and as the new House of Representatives had yet to convene, no Speaker either,[13] leaving no one at all in the line of succession after the vice president. When Congress convened in December 1885, President Cleveland asked for a revision of the 1792 act, which was passed in 1886. Congress replaced the President pro tempore and Speaker with officers of the President's Cabinet with the Secretary of State first in line. In the first 100 years of the United States, six former Secretaries of State had gone on to be elected President, while only two congressional leaders had advanced to that office. As a result, changing the order of the line of succession seemed reasonable.[citation needed]
Presidential Succession Act 1947[edit]
Wikisource has original text related to this article:
Presidential Succession Act 1947
The Presidential Succession Act of 1947, signed into law by President Harry S. Truman, added the Speaker of the House and President pro tempore back in the line, but switched the two from the 1792 order. It remains the sequence used today[update]. Since the reorganization of the military in 1947 had merged the War Department (which governed the Army) with the Department of the Navy into the Department of Defense, the Secretary of Defense took the place in the order of succession previously held by the Secretary of War. The office of Secretary of the Navy, which had existed as a Cabinet-level position since 1798, had become subordinate to the Secretary of Defense in the military reorganization, and so was dropped from the line of succession in the 1947 Succession Act.
Postmaster General removed 1971[edit]
Until 1971, the Postmaster General, the head of the Post Office Department, was a member of the Cabinet, initially the last in the presidential line of succession before new officers were added. Once the Post Office Department was re-organized into the United States Postal Service, a special agency independent of the executive branch, the Postmaster General ceased to be a member of the Cabinet and was thus removed from the line of succession.[14]
Secretary of Homeland Security added 2006[edit]
The United States Department of Homeland Security was created in 2002. On March 9, 2006, pursuant to the renewal of the Patriot Act as Pub.L. 109–177, the Secretary of Homeland Security was added to the line of succession. The order of Cabinet members in the line has always been the same as the order in which their respective departments were established. Despite custom, many in Congress had wanted the Secretary to be placed at number eight on the list – below the Attorney General, above the Secretary of the Interior, and in the position held by the Secretary of the Navy prior to the creation of the Secretary of Defense – because the Secretary, already in charge of disaster relief and security, would presumably be more prepared to take over the presidency than some of the other Cabinet secretaries.[citation needed] Despite this, the 2006 law explicitly specifies that the "Secretary of Homeland Security" follows the "Secretary of Veterans Affairs" in the succession, effectively at the end of the list.[15]
Successions beyond Vice President[edit]
David Rice Atchison's tombstone (Plattsburg, Missouri), including the claim "President of the United States for One Day"
While nine Vice Presidents have succeeded to the office upon the death or resignation of the President, and two Vice Presidents have temporarily served as acting President, no other officer has ever been called upon to act as President. On March 4, 1849, President Zachary Taylor's term began, but he declined to be sworn in on a Sunday, citing religious beliefs, and the Vice President was not sworn either. As the last President pro tempore of the Senate, David Rice Atchison was thought by some to be next in line after the Vice President, and his tombstone claims that he was US President for the day. However, Atchison took no oath of office to the presidency either, and his term as Senate President pro tempore had by then expired.[16][17]
In 1865, when Andrew Johnson assumed the Presidency on the death of Abraham Lincoln, the office of Vice President became vacant. At that time, the Senate President pro tempore was next in line to the presidency. In 1868, Johnson was impeached by the House of Representatives and subjected to trial in the Senate, and if he had been convicted and thereby removed from office, Senate President pro tempore Benjamin Wade would have become acting President. This posed a conflict of interest, as Wade's "guilty" vote could have been decisive in removing Johnson from office and giving himself presidential powers and duties. (Johnson was acquitted by a one-vote margin.)
In his book The Shadow Presidents, which he published in 1979, Michael Medved describes a situation that arose prior to the 1916 election, when the First World War was raging in Europe. In view of the contemporary international turmoil, President Woodrow Wilson thought that if he lost the election it would be better for his opponent to begin his administration straight away, instead of waiting through the lame duck period, which at that time had a duration of almost four months. President Wilson and his aides formed a plan to exploit the rule of succession so that his rival Charles Evans Hughes could take over the Presidency as soon as the result of the election was clear. The plan was that Wilson would appoint Hughes to the post of Secretary of State. Wilson and his Vice President Thomas R. Marshall would then resign, and as the Secretary of State was at that time designated next in line of succession, Hughes would become President immediately. As it happened, President Wilson won re-election, so the plan was never put into action.[18]
New President Gerald Ford is sworn in following the resignation of Richard Nixon on August 9, 1974.
Since the 25th Amendment's ratification, its Second Section, which addresses Vice Presidential succession as noted above, has been invoked twice.
During the 1973 Vice Presidential vacancy, House Speaker Carl Albert was first in line. As the Watergate scandal made President Nixon's removal or resignation possible, Albert would have become Acting President and—under Title 3, Section 19(c) of the U.S. Code—would have been able to "act as President until the expiration of the then current Presidential term" on January 20, 1977. Albert openly questioned whether it was appropriate for him, a Democrat, to assume the powers and duties of the presidency when there was a public mandate for the Presidency to be held by a Republican. Albert announced that should he need to assume the Presidential powers and duties, he would do so only as a caretaker. However, with the nomination and confirmation of Gerald Ford to the Vice Presidency, which marked the first time the Second Section of the Twenty-fifth Amendment was invoked, these series of events were never tested. Albert again became first-in-line during the first four months of Ford's Presidency, before the confirmation of Vice President Nelson Rockefeller,[citation needed] which marked the second time Section 2 of the Twenty-fifth Amendment was invoked.
In 1981, when President Ronald Reagan was shot, Vice President George H. W. Bush was traveling in Texas. Secretary of State Alexander Haig responded to a reporter's question regarding who was running the government by stating:
Constitutionally, gentlemen, you have the President, the Vice President and the Secretary of State in that order, and should the President decide he wants to transfer the helm to the Vice President, he will do so. He has not done that. As of now, I am in control here, in the White House, pending return of the Vice President and in close touch with him. If something came up, I would check with him, of course.[19]
A bitter dispute ensued over the meaning of Haig's remarks. Most people believed that Haig was referring to the line of succession and erroneously claimed to have temporary Presidential authority, due to his implied reference to the Constitution. Haig and his supporters, noting his familiarity with the line of succession from his time as White House Chief of Staff during Richard Nixon's resignation, said he only meant that he was the highest-ranking officer of the Executive branch on-site, managing things temporarily until the Vice President returned to Washington.[citation needed]
Constitutional concerns[edit]
Several constitutional law experts have raised questions as to the constitutionality of the provisions that the Speaker of the House and the President pro tempore of the Senate succeed to the presidency.[20] James Madison, one of the authors of the Constitution, raised similar constitutional questions about the Presidential Succession Act of 1792 in a 1792 letter to Edmund Pendleton.[21] Two of these issues can be summarized:
The term "Officer" in the relevant clause of the Constitution is most plausibly interpreted to mean an "Officer of the United States", who must be a member of the Executive or Judicial Branch. The Speaker and the President pro tempore are not officers in this sense.
Under the principle of separation of powers, the Constitution specifically disallows legislative officials from also serving in the executive branch. For the Speaker or the Senate President pro tempore to become Acting President, they must resign their position, at which point they are no longer in the line of succession. This is seen by some to form a constitutional paradox. However, the current Act specifies that the Speaker becomes President "upon his resignation as Speaker" and as a member of Congress,[22] allowing for a seamless transition.
In 2003 the Continuity of Government Commission suggested that the current law has "at least seven significant issues ... that warrant attention", specifically:[23]
The reality that all figures in the current line of succession work and reside in the vicinity of Washington, D.C. In the event of a nuclear, chemical, or biological attack, it is possible that everyone on the list would be killed or incapacitated.
Doubt (such as those expressed above by James Madison) that congressional leaders are eligible to act as President.
A concern about the wisdom of including the President pro tempore in the line of succession as the "largely honorific post traditionally held by the longest-serving Senator of the majority party". For example, from January 20, 2001, to June 6, 2001, the President pro tempore was then-98-year-old Strom Thurmond of South Carolina.
A concern that the current line of succession can force the presidency to abruptly switch parties mid-term, as the President, Speaker, and the President pro tempore are not necessarily of the same party as each other.
A concern that the succession line is ordered by the dates of creation of the various executive departments, without regard to the skills or capacities of the persons serving as their Secretary.
The fact that, should a Cabinet member begin to act as President, the law allows the House to elect a new Speaker (or the Senate, a new President pro tempore), who could in effect remove the Cabinet member and assume the office themselves at any time.
The absence of a provision where a President is disabled and the vice presidency is vacant (for example, if an assassination attempt simultaneously wounded the President and killed the Vice President).
See also[edit]
Central Locator System
Designated survivor
List of United States presidential assassination attempts and plots
United States presidential line of succession in fiction
References[edit]
Jump up ^ 3 U.S.C. § 19
Jump up ^ "Is the Presidential Succession Law Constitutional?", Akhil Reed Amar, Stanford Law Review, November 1995.
Jump up ^ First Report p. 4, continuityofgovernment.org Retrieved December 28, 2010 via Internet Archive to repair dead link
Jump up ^ Wilson, Reid (October 20, 2013). "The Presidential order of succession". The Washington Post. ISSN 0190-8286. Retrieved November 10, 2015.
Jump up ^ "The Continuity of the Presidency," Continuity of Government Commission, June 2009, p. 34. (Archived by WebCite at [1] Accessed: 2012-05-23)
Jump up ^ "Special Message to the Congress on the Succession to the Presidency. June 19, 1945" by President Harry S. Truman; from the American Presidency Project archives
^ Jump up to: a b Wildavsky, Aaron B. (2003). Horowitz, Irving, ed. The revolt against the masses and other essays on politics and public policy (Google Books). Transaction Publishers. p. 122. ISBN 978-0-7658-0960-5. Retrieved July 24, 2009.
Jump up ^ "kipnotes.com - Nachrichten zum trading -". kipnotes.com - Nachrichten zum trading.
^ Jump up to: a b c Schlesinger, Arthur M., Jr. (Autumn 1974). "On the Presidential Succession". Political Science Quarterly. 89 (3): 475, 495–496. JSTOR 2148451.
^ Jump up to: a b c Rankin, Robert S. (Feb 1946). "Presidential Succession in the United States". The Journal of Politics. 8 (1): 44–56. JSTOR 2125607. doi:10.2307/2125607.
^ Jump up to: a b c d e Abbott, Philip (Dec 2005). "Accidental Presidents: Death, Assassination, Resignation, and Democratic Succession". Presidential Studies Quarterly. 35 (4): 627, 638. JSTOR 27552721. doi:10.1111/j.1741-5705.2005.00269.x.
Jump up ^ Crapol, Edward P. (2006). John Tyler: the accidental president. UNC Press Books. p. 10. ISBN 978-0-8078-3041-3. OCLC 469686610.
Jump up ^ Lewis, Charlton Thomas; Willsey, Joseph H. (1895). Harper's Book of Facts: a Classified History of the World; Embracing Science, Literature, and Art. New York: Harper & Brothers. p. 884. LCCN 01020386. Retrieved April 24, 2011.
Jump up ^ "U.S. Postmasters General". postalmuseum.si.edu. Retrieved November 10, 2015.
Jump up ^ US Congress (2005). USA PATRIOT Improvement and Reauthorization Act OF 2005. 109th Congress Public Law 177. Retrieved on October 29, 2015 from http://www.gpo.gov/fdsys/pkg/PLAW-109publ177/html/PLAW-109publ177.htm.
Jump up ^ "1801: President for a Day -- March 4, 1849". May 29, 2014.
Jump up ^ David Mikkelson (October 8, 2007). "President for a Day". snopes.
Jump up ^ Medved, Michael (1979). The Shadow Presidents: The Secret History of the Chief Executives and Their Top Aides. New York, U.S.A.: Times Books. p. 149. ISBN 0-8129-0816-3.
Jump up ^ "Alexander Haig", Time, April 2, 1984
Jump up ^ "Is the Presidential Succession Law Constitutional?", Akhil Reed Amar, Stanford Law Review, November 1995.
Jump up ^ "Article 2, Section 1, Clause 6: James Madison to Edmund Pendleton".
Jump up ^ Wikisource:Presidential Succession Act 1947
Jump up ^ First Report p. 4, continuityofgovernment.org Retrieved December 28, 2010 via Internet Archive to repair dead link
Further reading[edit]
Ask Gleaves: Presidential Succession from the website of Grand Valley State University
3 U.S.C. § 19 – "Vacancy in offices of both President and Vice President; officers eligible to act"
Presidential Succession Act of 1792, 1 Stat. 239 (from the American Memory website of the Library of Congress
"Presidential Line of Succession Examined", a September 20, 2003 article from the Milwaukee Journal Sentinel
"WI Presidential Succession Act of 1947 held unconstitutional", David Tenner, Usenet group: soc.history.what-if, January 14, 2003.
Fools, Drunkards, & Presidential Succession from the Federalist Society website
Testimony of M. Miller Baker from the U.S. Senate website
External links[edit]
The Presidency: Preserving Our Institutions Second report of the Continuity of Government Commission, June 2009 (includes the text of the Presidential Succession Acts of 1792, 1886 and 1947)
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Jump up ^ Never confirmed by and with the advice and consent of the Senate and thus ineligible for the presidential line of succession.
Jump up ^ Although Elaine Chao is the current Secretary of Transportation, she is not a natural-born citizen (acquired U.S. citizenship by naturalization) and thus ineligible for the Presidency.
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