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Karrikin

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265:. Homologs of MAX2 are also required for strigolactone signaling in rice (known as DWARF3) petunia (DAD2) and pea (RMS4). Karrikin signaling also requires a protein named SUPPRESSOR OF MORE AXILARY GROWTH2-1 (SMAX1) which is a homolog of the DWARF53 protein required for strigolactone signaling in rice. SMAX1 and DWARF53 proteins could be involved in the control of cellular functions such as transport or transcription. The present model for karrikin and strigolactone signaling involves interaction of KAI2 or DWARF14 with SMAX1 or DWARF53 proteins respectively, which targets those proteins for 178:
are known as "fire-followers", they emerge grow quickly, flower and produce new seeds, which fall to the ground. These seeds can remain in the soil for decades, until the next fire produces fresh karrikins. Plants with this lifestyle are known as fire ephemerals. They thrive because the fire removes competing vegetation and provides nutrients and light for the emerging seedlings. Plants in many families respond to smoke and karrikins, suggesting that this response has evolved independently in different groups.
31: 80:. When plant material burns, these carbohydrates convert to karrikins. Burning plant products, such as straw, filter paper, cigarettes, and some sugars, can also produce karrikins. Seed germination activity can be generated within 30 minutes of heating plant material at 180 °C (356 °F). The 177:
occur largely in the ash at the site of the fire. Rains occurring after the fire wash the karrikins into the soil where dormant seeds reside. The karrikins and water can provide a 'wake-up call' for such seeds, triggering germination of the soil seed bank. The plants that depend on karrikins to grow
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Karrikins are released into the air upon the burning of plants. Subsequently, karrikins then get deposited on the soil surface and stimulate seed germination after rainfall. Since karrikins are released from smoke, they are released in huge quantities. Some plants which are known as "fire-followers"
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respond to karrikin signaling. Other studies have found that seed of ostensibly fire-adapted species do not display a sensitivity to karrikins. The difference between fire-followers and plants that respond to karrikins is their dependence on karrikins. Plants' response to karrikins is fundamental
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It has long been known that compounds released from smoke stimulate seed germination. To identify the active compounds that contribute to seed germination activity, smoke compounds were separated by liquid fractionation and were each tested for their effects on seed germination activity. Bioassays
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Karrikins are produced by wildfires but all seed plants contain KAI2 proteins, raising the question of the usual function of this protein. There is compelling evidence that plants contain an endogenous compound that is perceived by KAI2 to control seed germination and plant development, but this
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hormones which involves hydrolysis by their receptor protein DWARF14, an alpha/beta hydrolase related to KAI2. The question of whether karrikins act directly in plants is controversial. While some studies suggest that karrikins can bind directly to KAI2 protein, others do not support this. It is
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and destruction of proteins which are responsible for different aspect of plant growth, like the outgrowth of lateral shoots. This means that strigolactones, upon their interaction with D3 and D14; ubiquinate, and destroy proteins like DWARF53, which are responsible for the outgrowth of lateral
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The gene for KAI2 protein is present in lower plants including algae and mosses, whereas the DWARF14 protein evolved with seed plants, probably as a result of duplication of KAI2 followed by functional specialisation. Karrikin signaling could have evolved with seed plants as a result of the
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Karrikins easily dissolve in water, they are transparent, and have a melting point of 118–119 °C. However, they are unstable at very high temperatures and during common daylight, which means that they decay more rapidly than common active compounds which are not sensitive to sunlight.
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possible that karrikins produced by wildfires are converted to a different compound by the plant, before interaction with KAI2. The ability of different plants to carry out this conversion could partly explain differences in their ability to respond to karrikins and to smoke.
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are unable to germinate without karrikins. Fire-followers need rain after massive fires in order to germinate; this means that they may remain dormant and viable for decades until the right combination of fire occur in proper succession.
319:. Such responses could provide seedlings with an advantage as they emerge into the post-fire landscape. The KAI2 protein is also required for leaf development, implying that karrikins could influence other aspects of plant growth. 194:
because karrikins mimic the strigolactone hormones which are originally required for growth in plants. Fire-followers, on the other hand, have fine-tuned their responses according to the availability of karrikins. 
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mutants which failed to respond to karrikins. In rice, strigolactones interact with the F-box protein knowns as DWARF3 upon their hydrolysis by the DWARF14 (also known as D14-type proteins). This interaction targets the
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has been shown to respond to the two signals; KAR1, and KAR2. The two genes, MORE AXILLARY GROWTH2 (MAX2) and KARRIKIN-INSENSITIVE2 (KAI2) are essential for understanding the actions of karrikins and were discovered in
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Scaffidi A, Waters MT, Skelton BW, Bond CS, Sobolev AN, Bythell-Douglas R, et al. (May 2012). "Solar irradiation of the seed germination stimulant karrikinolide produces two novel head-to-head cage dimers".
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proteins known as KARRIKIN-INSENSITIVE1 (KAI1 or MAX2) in order to be able to interact with KARRIKIN-INSENSITIVE2 which is responsible for hypocotyl elongation and the inhibition of seed germination. The
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Gutjahr C, Gobbato E, Choi J, Riemann M, Johnston MG, Summers W, et al. (December 2015). "Rice perception of symbiotic arbuscular mycorrhizal fungi requires the karrikin receptor complex".
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van Staden J, Sparg SG, Kulkarni MG, Light ME (2006-08-01). "Post-germination effects of the smoke-derived compound 3-methyl-2H-furopyran-2-one, and its potential as a preconditioning agent".
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of KAI2, therefore stimulate seed germination and inhibits hypocotyl elongation. Karrikins could be used as agricultures, considering the environmental challenges that are occurring nowadays.
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of amino acids which is essential for activity, consistent with the hypothesis that KAI2 hydrolyses its ligand. This model is consistent with the perception of the chemically related
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Flematti GR, Scaffidi A, Dixon KW, Smith SM, Ghisalberti EL (February 2011). "Production of the seed germination stimulant karrikinolide from combustion of simple carbohydrates".
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Nelson DC, Flematti GR, Ghisalberti EL, Dixon KW, Smith SM (2012-01-01). "Regulation of seed germination and seedling growth by chemical signals from burning vegetation".
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Nelson DC, Flematti GR, Ghisalberti EL, Dixon KW, Smith SM (2012-06-02). "Regulation of seed germination and seedling growth by chemical signals from burning vegetation".
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Flematti GR, Ghisalberti EL, Dixon KW, Trengove RD (October 2009). "Identification of alkyl substituted 2H-furopyran-2-ones as germination stimulants present in smoke".
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The activity of karrikins requires an F-box protein named MORE AXILLARY GROWTH-2 (MAX2) in Arabidopsis. This protein is also required for strigolactone signaling in
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Chiwocha SD, Dixon KW, Flematti GR, Ghisalberti EL, Merritt DJ, Nelson DC, et al. (2009-10-01). "Karrikins: A new family of plant growth regulators in smoke".
1042:"A Selaginella moellendorffii Ortholog of KARRIKIN INSENSITIVE2 Functions in Arabidopsis Development but Cannot Mediate Responses to Karrikins or Strigolactones" 202:
Carbon, hydrogen, and oxygen make up the two ring structures found in karrikins, one of which is a six-membered, heterocyclic ring with a molecular formula of C
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found in the smoke of burning plant material. Karrikins help stimulate seed germination and plant development because they mimic a signaling hormone known as
64:) was shown to be responsible for this effect. Later, several closely related compounds were discovered in smoke, and are collectively known as karrikins. 1687: 998:"Substrate-Induced Degradation of the α/β-Fold Hydrolase KARRIKIN INSENSITIVE2 Requires a Functional Catalytic Triad but Is Independent of MAX2" 161:, Flematti proposed changing the name of the molecule and its related compounds to karrikin. One of the first recorded Western Australian 773: 496:"Ecological relevance of strigolactones in nutrient uptake and other abiotic stresses, and in plant-microbe interactions below-ground" 834:"Karrikins discovered in smoke trigger Arabidopsis seed germination by a mechanism requiring gibberellic acid synthesis and light" 88:
sugar. There is no evidence that karrikins occur naturally in plants, but it has been postulated that karrikin-like molecules do.
1816: 957:"Specialisation within the DWARF14 protein family confers distinct responses to karrikins and strigolactones in Arabidopsis" 181:
Fire-followers are not the only plants that respond to karrikins. Seeds from a number of different flowering families like
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Smoke from wildfires or bushfires has been known for a long time to stimulate the germination of seeds. In 2004, the
1493:"Karrikins: Regulators Involved in Phytohormone Signaling Networks during Seed Germination and Seedling Development" 157:, was initially named gavinone in reference to its discovery by chemist Gavin Flematti. After consulting with an 1806: 1673: 881:
George N (August 2009). "Does karrikinolide improve the germination and seedling vigour of switchgrass?".
174: 1244:"SUPPRESSOR OF MORE AXILLARY GROWTH2 1 controls seed germination and seedling development in Arabidopsis" 1765: 50: 832:
Nelson DC, Riseborough JA, Flematti GR, Stevens J, Ghisalberti EL, Dixon KW, Smith SM (February 2009).
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Bennett T, Leyser O (December 2014). "Strigolactone signalling: standing on the shoulders of DWARFs".
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requires an alpha/beta-fold hydrolase named KARRIKIN-INSENSITIVE-2 (KAI2). The KAI2 protein has a
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Karrikins not only stimulate seed germination, but are reported to increase seedling vigour. In
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Smith SM, Li J (October 2014). "Signalling and responses to strigolactones and karrikins".
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The mode of action of karrikins has been largely determined using the genetic resources of
1717: 1442:"Q&A: What are strigolactones and why are they important to plants and soil microbes?" 743: 567: 245: 73: 1183:
Nelson DC, Scaffidi A, Dun EA, Waters MT, Flematti GR, Dixon KW, et al. (May 2011).
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Karrikins are formed by the heating or combustion of carbohydrates, including sugars and
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Waters MT, Nelson DC, Scaffidi A, Flematti GR, Sun YK, Dixon KW, Smith SM (April 2012).
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in the soil, which enhances plant growth and leads to an increase in plant branching.
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Proceedings of the National Academy of Sciences of the United States of America
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Proceedings of the National Academy of Sciences of the United States of America
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Six karrikins have so far been discovered in smoke, and they are designated KAR
1606: 1418: 1126:"Smoke-derived karrikin perception by the α/β-hydrolase KAI2 from Arabidopsis" 1102: 894: 540: 512: 495: 494:
Andreo-Jimenez B, Ruyter-Spira C, Bouwmeester HJ, Lopez-Raez JA (2015-09-01).
405: 329: 219: 57: 1350:"D14-SCF(D3)-dependent degradation of D53 regulates strigolactone signalling" 1291:
Jiang L, Liu X, Xiong G, Liu H, Chen F, Wang L, et al. (December 2013).
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Waters MT, Scaffidi A, Moulin SL, Sun YK, Flematti GR, Smith SM (July 2015).
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Zhou F, Lin Q, Zhu L, Ren Y, Zhou K, Shabek N, et al. (December 2013).
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is also known as karrikinolode and was the first karrikin to be discovered.
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shoots, and for the inhibition of stem thickening and root branching. In
85: 17: 1544:"Potential of Karrikins as Novel Plant Growth Regulators in Agriculture" 1373: 1316: 1647: 973: 956: 930: 215: 186: 810: 654: 30: 182: 49:. Strigolactones are hormones that help increase growth of symbiotic 1491:
Meng Y, Shuai H, Luo X, Chen F, Zhou W, Yang W, Shu K (2017-01-24).
1712: 1293:"DWARF 53 acts as a repressor of strigolactone signalling in rice" 592:
Flematti GR, Ghisalberti EL, Dixon KW, Trengove RD (August 2004).
290:, karrikins work in a similar way to strigolactones; they require 211: 81: 29: 1665: 165:
words for 'smoke' from the Perth area in the 1830s, is 'karrik'.
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identified several related compounds that were named karrikins.
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He T, Pausas JG, Belcher CM, Schwilk DW, Lamont BB (May 2012).
389:"What are karrikins and how were they 'discovered' by plants?" 328:
divergence of KAI2 and DWARF14 functions, possibly during the
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Chemical structures of karrikins (KAR1, KAR2, KAR3, and KAR4)
1622:"Fire-adapted traits of Pinus arose in the fiery Cretaceous" 1242:
Stanga JP, Smith SM, Briggs WR, Nelson DC (September 2013).
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Guo Y, Zheng Z, La Clair JJ, Chory J, Noel JP (May 2013).
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Antala M, Sytar O, Rastogi A, Brestic M (December 2019).
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Waters MT, Scaffidi A, Flematti G, Smith SM (May 2015).
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moiety of karrikins is probably directly derived from a
594:"A compound from smoke that promotes seed germination" 766:
Chemistry : human activity, chemical reactivity
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compound is neither a karrikin nor a strigolactone.
471:"Strigolactone - Latest research and news | Nature" 387:Flematti GR, Dixon KW, Smith SM (December 2015). 153:The first karrikin discovered, abbreviated as KAR 382: 380: 378: 376: 374: 587: 585: 438: 436: 434: 372: 370: 368: 366: 364: 362: 360: 358: 356: 354: 1035: 1033: 950: 948: 725: 723: 1681: 315:, resulting in shorter hypocotyls and larger 8: 1093:. SI: Cell signalling and gene regulation. 1688: 1674: 1666: 799:Journal of Agricultural and Food Chemistry 643:Journal of Agricultural and Food Chemistry 1637: 1569: 1559: 1518: 1508: 1467: 1457: 1381: 1324: 1267: 1218: 1208: 1159: 1149: 1065: 1013: 972: 857: 609: 511: 414: 404: 332:period when fires were common on Earth. 198:Structure and physicochemical properties 350: 744:10.1146/annurev-arplant-042811-105545 568:10.1146/annurev-arplant-042811-105545 7: 919:Organic & Biomolecular Chemistry 527:Halford B (2010). "Smoke Signals". 214:, and the other is a five-membered 132:are the most active karrikins. KAR 25: 1639:10.1111/j.1469-8137.2012.04079.x 1407:Current Opinion in Plant Biology 1091:Current Opinion in Plant Biology 529:Chemical & Engineering News 311:, karrikins influence seedling 732:Annual Review of Plant Biology 556:Annual Review of Plant Biology 457:10.1016/j.plantsci.2009.06.007 1: 240:. Perception of karrikins by 51:arbuscular mycorrhizal fungi 883:Seed Science and Technology 1833: 1497:Frontiers in Plant Science 1015:10.1016/j.molp.2014.12.020 1741: 1703: 1607:10.1016/j.fcr.2005.12.007 1419:10.1016/j.pbi.2014.08.001 1103:10.1016/j.pbi.2014.06.003 895:10.15258/sst.2009.37.1.30 541:10.1021/cen-v088n015.p037 513:10.1007/s11104-015-2544-z 406:10.1186/s12915-015-0219-0 169:The response to karrikins 27:A plant growth regulator 1817:Plant growth regulators 1510:10.3389/fpls.2016.02021 1459:10.1186/1741-7007-12-19 1440:Smith SM (March 2014). 1210:10.1073/pnas.1100987108 1151:10.1073/pnas.1306265110 768:. : Brooks Cole. 2014. 698:10.1126/science.aac9715 611:10.1126/science.1099944 303:Effects on plant growth 43:plant growth regulators 1766:Plant peptide hormones 173:Karrikins produced by 35: 1561:10.3390/plants9010043 1260:10.1104/pp.113.221259 850:10.1104/pp.108.131516 336:Response to wildfires 33: 1595:Field Crops Research 1409:. 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Index

Karrikins

plant growth regulators
strigolactone
arbuscular mycorrhizal fungi
butenolide
polysaccharides
cellulose
pyran
pyranose
etymologist
Noongar
bushfires
tomatoes
lettuce
trees
pyran
lactone
butenolide
Arabidopsis thaliana
catalytic triad
strigolactone
ubiquitination
ubiquitination
homologous
ubiquination
photomorphogenesis
cotyledons
Cretaceous

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