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Vernalization

February 15, 2024

Vernalization (from Latin vernus 'of the spring') is the induction of a plant's flowering process by exposure to the prolonged cold of winter, or by an artificial equivalent. After vernalization, plants have acquired the ability to flower, but they may require additional seasonal cues or weeks of growth before they will actually do so. The term is sometimes used to refer to the need of herbal (non-woody) plants for a period of cold dormancy in order to produce new shoots and leaves,[1] but this usage is discouraged.[2]

Many species of henbane require vernalization before flowering.

Many plants grown in temperate climates require vernalization and must experience a period of low winter temperature to initiate or accelerate the flowering process. This ensures that reproductive development and seed production occurs in spring and winters, rather than in autumn.[3] The needed cold is often expressed in chill hours. Typical vernalization temperatures are between 1 and 7 degrees Celsius (34 and 45 degrees Fahrenheit).[4]

For many perennial plants, such as fruit tree species, a period of cold is needed first to induce dormancy and then later, after the requisite period of time, re-emerge from that dormancy prior to flowering. Many monocarpic winter annuals and biennials, including some ecotypes of Arabidopsis thaliana[5] and winter cereals such as wheat, must go through a prolonged period of cold before flowering occurs.

History of vernalization research edit

In the history of agriculture, farmers observed a traditional distinction between "winter cereals", whose seeds require chilling (to trigger their subsequent emergence and growth), and "spring cereals", whose seeds can be sown in spring, and germinate, and then flower soon thereafter. Scientists in the early 19th century had discussed how some plants needed cold temperatures to flower. In 1857 an American agriculturist John Hancock Klippart, Secretary of the Ohio Board of Agriculture, reported the importance and effect of winter temperature on the germination of wheat. One of the most significant works was by a German plant physiologist Gustav Gassner who made a detailed discussion in his 1918 paper. Gassner was the first to systematically differentiate the specific requirements of winter plants from those of summer plants, and also that early swollen germinating seeds of winter cereals are sensitive to cold.[6]

In 1928, the Soviet agronomist Trofim Lysenko published his works on the effects of cold on cereal seeds, and coined the term "яровизация" ("jarovization") to describe a chilling process he used to make the seeds of winter cereals behave like spring cereals (Jarovoe in Russian, originally from jar meaning fire or the god of spring). Lysenko himself translated the term into "vernalization" (from the Latin vernum meaning Spring). After Lysenko the term was used to explain the ability of flowering in some plants after a period of chilling due to physiological changes and external factors. The formal definition was given in 1960 by a French botanist P. Chouard, as "the acquisition or acceleration of the ability to flower by a chilling treatment".[7]

Lysenko's 1928 paper on vernalization and plant physiology drew wide attention due to its practical consequences for Russian agriculture. Severe cold and lack of winter snow had destroyed many early winter wheat seedlings. By treating wheat seeds with moisture as well as cold, Lysenko induced them to bear a crop when planted in spring.[8] Later however, according to Richard Amasino, Lysenko inaccurately asserted that the vernalized state could be inherited, i.e. the offspring of a vernalized plant would behave as if they themselves had also been vernalized and would not require vernalization in order to flower quickly.[9] Opposing this view and supporting Lysenko's claim, Xiuju Li and Yongsheng Liu have detailed experimental evidence from the USSR, Hungary, Bulgaria and China that shows the conversion between spring wheat and winter wheat, positing that "it is not unreasonable to postulate epigenetic mechanisms that could plausibly result in the conversion of spring to winter wheat or vice versa."[10]

Early research on vernalization focused on plant physiology; the increasing availability of molecular biology has made it possible to unravel its underlying mechanisms.[9] For example, a lengthening daylight period (longer days), as well as cold temperatures are required for winter wheat plants to go from the vegetative to the reproductive state; the three interacting genes are called VRN1, VRN2, and FT (VRN3).[11]

In Arabidopsis thaliana edit

 
Arabidopsis thaliana rosette before vernalization, with no floral spike

Arabidopsis thaliana ("thale cress") is a much-studied model for vernalization. Some ecotypes (varieties), called "winter annuals", have delayed flowering without vernalization; others ("summer annuals") do not.[12][self-published source?] The genes that underlie this difference in plant physiology have been intensively studied.[9]

The reproductive phase change of A. thaliana occurs by a sequence of two related events: first, the bolting transition (flower stalk elongates), then the floral transition (first flower appears).[13] Bolting is a robust predictor of flower formation, and hence a good indicator for vernalization research.[13]

In winter annual Arabidopsis, vernalization of the meristem appears to confer competence to respond to floral inductive signals. A vernalized meristem retains competence for as long as 300 days in the absence of an inductive signal.[12]

At the molecular level, flowering is repressed by the protein Flowering Locus C (FLC), which binds to and represses genes that promote flowering, thus blocking flowering.[3][14] Winter annual ecotypes of Arabidopsis have an active copy of the gene FRIGIDA (FRI), which promotes FLC expression, thus repression of flowering.[15] Prolonged exposure to cold (vernalization) induces expression of VERNALIZATION INSENSTIVE3, which interacts with the VERNALIZATION2 (VRN2) polycomb-like complex to reduce FLC expression through chromatin remodeling.[16] Levels of VRN2 protein increase during long-term cold exposure as a result of inhibition of VRN2 turnover via its N-degron.[17] The events of histone deacetylation at Lysine 9 and 14 followed by methylation at Lys 9 and 27 is associated with the vernalization response. The epigenetic silencing of FLC by chromatin remodeling is also thought to involve the cold-induced expression of antisense FLC COOLAIR[18][19] or COLDAIR transcripts.[20] Vernalization is registered by the plant by the stable silencing of individual FLC loci.[21] The removal of silent chromatin marks at FLC during embryogenesis prevents the inheritance of the vernalized state.[22]

Since vernalization also occurs in flc mutants (lacking FLC), vernalization must also activate a non-FLC pathway.[23][self-published source?] A day-length mechanism is also important.[11] Vernalization response works in concert with the photo-periodic genes CO, FT, PHYA, CRY2 to induce flowering.

Devernalization edit

It is possible to devernalize a plant by exposure to sometimes low and high temperatures subsequent to vernalization. For example, commercial onion growers store sets at low temperatures, but devernalize them before planting, because they want the plant's energy to go into enlarging its bulb (underground stem), not making flowers.[24]

See also edit

References edit

  1. ^ Sokolski, K.; Dovholuk, A.; Dovholuk, L.; Faletra, P. (1997). "Axenic seed culture and micropropagation of Cypripedium reginae". Selbyana. 18 (2): 172–82. JSTOR 41760430.
  2. ^ Chouard, P. (June 1960). "Vernalization and its relations to dormancy". Annual Review of Plant Physiology. Annual Reviews. 11: 191–238. doi:10.1146/annurev.pp.11.060160.001203.
  3. ^ a b Sung, Sibum; He, Yuehui; Eshoo, Tifani W; Tamada, Yosuke; Johnson, Lianna; Nakahigashi, Kenji; Goto, Koji; Jacobsen, Steve E; Amasino, Richard M (2006). "Epigenetic maintenance of the vernalized state in Arabidopsis thaliana requires LIKE HETEROCHROMATIN PROTEIN 1". Nature Genetics. 38 (6): 706–10. doi:10.1038/ng1795. PMID 16682972. S2CID 2855447.
  4. ^ Taiz, Lincoln; Murphy, Angus (2015). Plant Physiology and Development. Sunderland, Massachusetts (USA): Sinauer Associates. p. 605. ISBN 978-1-60535-255-8.
  5. ^ Michaels, Scott D.; He, Yuehui; Scortecci, Katia C.; Amasino, Richard M. (2003). "Attenuation of FLOWERING LOCUS C activity as a mechanism for the evolution of summer-annual flowering behavior in Arabidopsis". Proceedings of the National Academy of Sciences. 100 (17): 10102–7. Bibcode:2003PNAS..10010102M. doi:10.1073/pnas.1531467100. JSTOR 3147669. PMC 187779. PMID 12904584.
  6. ^ Chouard, P. (1960). "Vernalization and its relations to dormancy". Annual Review of Plant Physiology. 11 (1): 191–238. doi:10.1146/annurev.pp.11.060160.001203.
  7. ^ Poltronieri, Palmiro; Hong, Yiguo (2015). Applied Plant Genomics and Biotechnology. Cambridge (UK): Woodhead Publishing. p. 121. ISBN 978-0-08-100068-7.
  8. ^ Roll-Hansen, Nils (1985). "A new perspective on Lysenko?". Annals of Science. Taylor & Francis. 42 (3): 261–278. doi:10.1080/00033798500200201. PMID 11620694.
  9. ^ a b c Amasino, R. (2004). "Vernalization, competence, and the epigenetic memory of winter". The Plant Cell. 16 (10): 2553–2559. doi:10.1105/tpc.104.161070. PMC 520954. PMID 15466409.
  10. ^ Li, Xiuju; Liu, Yongsheng (6 May 2010). "The conversion of spring wheat into winter wheat and vice versa: false claim or Lamarckian inheritance?". Journal of Biosciences. 35 (2): 321–325. doi:10.1007/s12038-010-0035-1. ISSN 0250-5991. PMID 20689187. S2CID 10527354.
  11. ^ a b Trevaskis, Ben; Hemming, Megan N.; Dennis, Elizabeth S. (August 2007). "The molecular basis of vernalization-induced flowering in cereals". Trends in Plant Science. Elsevier. 12 (8): 352–357. doi:10.1016/j.tplants.2007.06.010. PMID 17629542.
  12. ^ a b "Vernalisation response". Plant Biology. Retrieved 26 January 2011.[self-published source]
  13. ^ a b Pouteau, Sylvie; Albertini, Catherine (2009). "The significance of bolting and floral transitions as indicators of reproductive phase change in Arabidopsis". Journal of Experimental Botany. 60 (12): 3367–77. doi:10.1093/jxb/erp173. PMID 19502535.
  14. ^ Amasino, Richard (2010). "Seasonal and developmental timing of flowering". The Plant Journal. 61 (6): 1001–13. doi:10.1111/j.1365-313X.2010.04148.x. PMID 20409274.
  15. ^ Choi, Kyuha; Kim, Juhyun; Hwang, Hyun-Ju; Kim, Sanghee; Park, Chulmin; Kim, Sang Yeol; Lee, Ilha (2011). "The FRIGIDA Complex Activates Transcription ofFLC, a Strong Flowering Repressor in Arabidopsis, by Recruiting Chromatin Modification Factors". The Plant Cell. 23 (1): 289–303. doi:10.1105/tpc.110.075911. PMC 3051252. PMID 21282526.
  16. ^ Sung, Sibum; Amasino, Richard M. (2004). "Vernalization in Arabidopsis thaliana is mediated by the PHD finger protein VIN3". Nature. 427 (6970): 159–163. Bibcode:2004Natur.427..159S. doi:10.1038/nature02195. PMID 14712276. S2CID 4418494.
  17. ^ Gibbs, DJ; Tedds, HM; Labandera, AM; Bailey, M; White, MD; Hartman, S; Sprigg, C; Mogg, SL; Osborne, R; Dambire, C; Boeckx, T; Paling, Z; Voesenek, LACJ; Flashman, E; Holdsworth, MJ (21 December 2018). "Oxygen-dependent proteolysis regulates the stability of angiosperm polycomb repressive complex 2 subunit VERNALIZATION 2". Nature Communications. 9 (1): 5438. Bibcode:2018NatCo...9.5438G. doi:10.1038/s41467-018-07875-7. PMC 6303374. PMID 30575749.
  18. ^ http://www.jic.ac.uk/news/2014/10/plants-require-coolair-flower-spring 23 April 2015 at the Wayback Machine[full citation needed]
  19. ^ Csorba, Tibor; Questa, Julia I.; Sun, Qianwen; Dean, Caroline (2014). "Antisense COOLAIR mediates the coordinated switching of chromatin states atFLCduring vernalization". Proceedings of the National Academy of Sciences. 111 (45): 16160–5. Bibcode:2014PNAS..11116160C. doi:10.1073/pnas.1419030111. PMC 4234544. PMID 25349421.
  20. ^ Heo, J. B.; Sung, S. (2011). "Vernalization-Mediated Epigenetic Silencing by a Long Intronic Noncoding RNA". Science. 331 (6013): 76–9. Bibcode:2011Sci...331...76H. doi:10.1126/science.1197349. PMID 21127216. S2CID 19127414.
  21. ^ Angel, Andrew; Song, Jie; Dean, Caroline; Howard, Martin (2011). "A Polycomb-based switch underlying quantitative epigenetic memory". Nature. 476 (7358): 105–8. doi:10.1038/nature10241. PMID 21785438. S2CID 205225603.
  22. ^ Crevillén, Pedro; Yang, Hongchun; Cui, Xia; Greeff, Christiaan; Trick, Martin; Qiu, Qi; Cao, Xiaofeng; Dean, Caroline (2014). "Epigenetic reprogramming that prevents trans-generational inheritance of the vernalized state". Nature. 515 (7528): 587–90. Bibcode:2014Natur.515..587C. doi:10.1038/nature13722. PMC 4247276. PMID 25219852.
  23. ^ "Vernalisation pathway". Plant Biology. Retrieved 26 January 2011.[self-published source]
  24. ^ "Vernalization". Encyclopædia Britannica Online. Retrieved 3 September 2023. Devernalization can be brought about by high temperatures ... Onion sets ... are ... ready to flower ... temperatures above 26.7 °C (80 °F) ..., however, shifts the sets to the desired bulb-forming phase.

External links edit

February 15, 2024
vernalization, from, latin, vernus, spring, induction, plant, flowering, process, exposure, prolonged, cold, winter, artificial, equivalent, after, vernalization, plants, have, acquired, ability, flower, they, require, additional, seasonal, cues, weeks, growth. Vernalization from Latin vernus of the spring is the induction of a plant s flowering process by exposure to the prolonged cold of winter or by an artificial equivalent After vernalization plants have acquired the ability to flower but they may require additional seasonal cues or weeks of growth before they will actually do so The term is sometimes used to refer to the need of herbal non woody plants for a period of cold dormancy in order to produce new shoots and leaves 1 but this usage is discouraged 2 Many species of henbane require vernalization before flowering Many plants grown in temperate climates require vernalization and must experience a period of low winter temperature to initiate or accelerate the flowering process This ensures that reproductive development and seed production occurs in spring and winters rather than in autumn 3 The needed cold is often expressed in chill hours Typical vernalization temperatures are between 1 and 7 degrees Celsius 34 and 45 degrees Fahrenheit 4 For many perennial plants such as fruit tree species a period of cold is needed first to induce dormancy and then later after the requisite period of time re emerge from that dormancy prior to flowering Many monocarpic winter annuals and biennials including some ecotypes of Arabidopsis thaliana 5 and winter cereals such as wheat must go through a prolonged period of cold before flowering occurs Contents 1 History of vernalization research 2 In Arabidopsis thaliana 3 Devernalization 4 See also 5 References 6 External linksHistory of vernalization research editIn the history of agriculture farmers observed a traditional distinction between winter cereals whose seeds require chilling to trigger their subsequent emergence and growth and spring cereals whose seeds can be sown in spring and germinate and then flower soon thereafter Scientists in the early 19th century had discussed how some plants needed cold temperatures to flower In 1857 an American agriculturist John Hancock Klippart Secretary of the Ohio Board of Agriculture reported the importance and effect of winter temperature on the germination of wheat One of the most significant works was by a German plant physiologist Gustav Gassner who made a detailed discussion in his 1918 paper Gassner was the first to systematically differentiate the specific requirements of winter plants from those of summer plants and also that early swollen germinating seeds of winter cereals are sensitive to cold 6 In 1928 the Soviet agronomist Trofim Lysenko published his works on the effects of cold on cereal seeds and coined the term yarovizaciya jarovization to describe a chilling process he used to make the seeds of winter cereals behave like spring cereals Jarovoe in Russian originally from jar meaning fire or the god of spring Lysenko himself translated the term into vernalization from the Latin vernum meaning Spring After Lysenko the term was used to explain the ability of flowering in some plants after a period of chilling due to physiological changes and external factors The formal definition was given in 1960 by a French botanist P Chouard as the acquisition or acceleration of the ability to flower by a chilling treatment 7 Lysenko s 1928 paper on vernalization and plant physiology drew wide attention due to its practical consequences for Russian agriculture Severe cold and lack of winter snow had destroyed many early winter wheat seedlings By treating wheat seeds with moisture as well as cold Lysenko induced them to bear a crop when planted in spring 8 Later however according to Richard Amasino Lysenko inaccurately asserted that the vernalized state could be inherited i e the offspring of a vernalized plant would behave as if they themselves had also been vernalized and would not require vernalization in order to flower quickly 9 Opposing this view and supporting Lysenko s claim Xiuju Li and Yongsheng Liu have detailed experimental evidence from the USSR Hungary Bulgaria and China that shows the conversion between spring wheat and winter wheat positing that it is not unreasonable to postulate epigenetic mechanisms that could plausibly result in the conversion of spring to winter wheat or vice versa 10 Early research on vernalization focused on plant physiology the increasing availability of molecular biology has made it possible to unravel its underlying mechanisms 9 For example a lengthening daylight period longer days as well as cold temperatures are required for winter wheat plants to go from the vegetative to the reproductive state the three interacting genes are called VRN1 VRN2 and FT VRN3 11 In Arabidopsis thaliana edit nbsp Arabidopsis thaliana rosette before vernalization with no floral spikeArabidopsis thaliana thale cress is a much studied model for vernalization Some ecotypes varieties called winter annuals have delayed flowering without vernalization others summer annuals do not 12 self published source The genes that underlie this difference in plant physiology have been intensively studied 9 The reproductive phase change of A thaliana occurs by a sequence of two related events first the bolting transition flower stalk elongates then the floral transition first flower appears 13 Bolting is a robust predictor of flower formation and hence a good indicator for vernalization research 13 In winter annual Arabidopsis vernalization of the meristem appears to confer competence to respond to floral inductive signals A vernalized meristem retains competence for as long as 300 days in the absence of an inductive signal 12 At the molecular level flowering is repressed by the protein Flowering Locus C FLC which binds to and represses genes that promote flowering thus blocking flowering 3 14 Winter annual ecotypes of Arabidopsis have an active copy of the gene FRIGIDA FRI which promotes FLC expression thus repression of flowering 15 Prolonged exposure to cold vernalization induces expression of VERNALIZATION INSENSTIVE3 which interacts with the VERNALIZATION2 VRN2 polycomb like complex to reduce FLC expression through chromatin remodeling 16 Levels of VRN2 protein increase during long term cold exposure as a result of inhibition of VRN2 turnover via its N degron 17 The events of histone deacetylation at Lysine 9 and 14 followed by methylation at Lys 9 and 27 is associated with the vernalization response The epigenetic silencing of FLC by chromatin remodeling is also thought to involve the cold induced expression of antisense FLC COOLAIR 18 19 or COLDAIR transcripts 20 Vernalization is registered by the plant by the stable silencing of individual FLC loci 21 The removal of silent chromatin marks at FLC during embryogenesis prevents the inheritance of the vernalized state 22 Since vernalization also occurs in flc mutants lacking FLC vernalization must also activate a non FLC pathway 23 self published source A day length mechanism is also important 11 Vernalization response works in concert with the photo periodic genes CO FT PHYA CRY2 to induce flowering Devernalization editIt is possible to devernalize a plant by exposure to sometimes low and high temperatures subsequent to vernalization For example commercial onion growers store sets at low temperatures but devernalize them before planting because they want the plant s energy to go into enlarging its bulb underground stem not making flowers 24 See also editStratification seeds References edit Sokolski K Dovholuk A Dovholuk L Faletra P 1997 Axenic seed culture and micropropagation of Cypripedium reginae Selbyana 18 2 172 82 JSTOR 41760430 Chouard P June 1960 Vernalization and its relations to dormancy Annual Review of Plant Physiology Annual Reviews 11 191 238 doi 10 1146 annurev pp 11 060160 001203 a b Sung Sibum He Yuehui Eshoo Tifani W Tamada Yosuke Johnson Lianna Nakahigashi Kenji Goto Koji Jacobsen Steve E Amasino Richard M 2006 Epigenetic maintenance of the vernalized state in Arabidopsis thaliana requires LIKE HETEROCHROMATIN PROTEIN 1 Nature Genetics 38 6 706 10 doi 10 1038 ng1795 PMID 16682972 S2CID 2855447 Taiz Lincoln Murphy Angus 2015 Plant Physiology and Development Sunderland Massachusetts USA Sinauer Associates p 605 ISBN 978 1 60535 255 8 Michaels Scott D He Yuehui Scortecci Katia C Amasino Richard M 2003 Attenuation of FLOWERING LOCUS C activity as a mechanism for the evolution of summer annual flowering behavior in Arabidopsis Proceedings of the National Academy of Sciences 100 17 10102 7 Bibcode 2003PNAS 10010102M doi 10 1073 pnas 1531467100 JSTOR 3147669 PMC 187779 PMID 12904584 Chouard P 1960 Vernalization and its relations to dormancy Annual Review of Plant Physiology 11 1 191 238 doi 10 1146 annurev pp 11 060160 001203 Poltronieri Palmiro Hong Yiguo 2015 Applied Plant Genomics and Biotechnology Cambridge UK Woodhead Publishing p 121 ISBN 978 0 08 100068 7 Roll Hansen Nils 1985 A new perspective on Lysenko Annals of Science Taylor amp Francis 42 3 261 278 doi 10 1080 00033798500200201 PMID 11620694 a b c Amasino R 2004 Vernalization competence and the epigenetic memory of winter The Plant Cell 16 10 2553 2559 doi 10 1105 tpc 104 161070 PMC 520954 PMID 15466409 Li Xiuju Liu Yongsheng 6 May 2010 The conversion of spring wheat into winter wheat and vice versa false claim or Lamarckian inheritance Journal of Biosciences 35 2 321 325 doi 10 1007 s12038 010 0035 1 ISSN 0250 5991 PMID 20689187 S2CID 10527354 a b Trevaskis Ben Hemming Megan N Dennis Elizabeth S August 2007 The molecular basis of vernalization induced flowering in cereals Trends in Plant Science Elsevier 12 8 352 357 doi 10 1016 j tplants 2007 06 010 PMID 17629542 a b Vernalisation response Plant Biology Retrieved 26 January 2011 self published source a b Pouteau Sylvie Albertini Catherine 2009 The significance of bolting and floral transitions as indicators of reproductive phase change in Arabidopsis Journal of Experimental Botany 60 12 3367 77 doi 10 1093 jxb erp173 PMID 19502535 Amasino Richard 2010 Seasonal and developmental timing of flowering The Plant Journal 61 6 1001 13 doi 10 1111 j 1365 313X 2010 04148 x PMID 20409274 Choi Kyuha Kim Juhyun Hwang Hyun Ju Kim Sanghee Park Chulmin Kim Sang Yeol Lee Ilha 2011 The FRIGIDA Complex Activates Transcription ofFLC a Strong Flowering Repressor in Arabidopsis by Recruiting Chromatin Modification Factors The Plant Cell 23 1 289 303 doi 10 1105 tpc 110 075911 PMC 3051252 PMID 21282526 Sung Sibum Amasino Richard M 2004 Vernalization in Arabidopsis thaliana is mediated by the PHD finger protein VIN3 Nature 427 6970 159 163 Bibcode 2004Natur 427 159S doi 10 1038 nature02195 PMID 14712276 S2CID 4418494 Gibbs DJ Tedds HM Labandera AM Bailey M White MD Hartman S Sprigg C Mogg SL Osborne R Dambire C Boeckx T Paling Z Voesenek LACJ Flashman E Holdsworth MJ 21 December 2018 Oxygen dependent proteolysis regulates the stability of angiosperm polycomb repressive complex 2 subunit VERNALIZATION 2 Nature Communications 9 1 5438 Bibcode 2018NatCo 9 5438G doi 10 1038 s41467 018 07875 7 PMC 6303374 PMID 30575749 http www jic ac uk news 2014 10 plants require coolair flower spring Archived 23 April 2015 at the Wayback Machine full citation needed Csorba Tibor Questa Julia I Sun Qianwen Dean Caroline 2014 Antisense COOLAIR mediates the coordinated switching of chromatin states atFLCduring vernalization Proceedings of the National Academy of Sciences 111 45 16160 5 Bibcode 2014PNAS 11116160C doi 10 1073 pnas 1419030111 PMC 4234544 PMID 25349421 Heo J B Sung S 2011 Vernalization Mediated Epigenetic Silencing by a Long Intronic Noncoding RNA Science 331 6013 76 9 Bibcode 2011Sci 331 76H doi 10 1126 science 1197349 PMID 21127216 S2CID 19127414 Angel Andrew Song Jie Dean Caroline Howard Martin 2011 A Polycomb based switch underlying quantitative epigenetic memory Nature 476 7358 105 8 doi 10 1038 nature10241 PMID 21785438 S2CID 205225603 Crevillen Pedro Yang Hongchun Cui Xia Greeff Christiaan Trick Martin Qiu Qi Cao Xiaofeng Dean Caroline 2014 Epigenetic reprogramming that prevents trans generational inheritance of the vernalized state Nature 515 7528 587 90 Bibcode 2014Natur 515 587C doi 10 1038 nature13722 PMC 4247276 PMID 25219852 Vernalisation pathway Plant Biology Retrieved 26 January 2011 self published source Vernalization Encyclopaedia Britannica Online Retrieved 3 September 2023 Devernalization can be brought about by high temperatures Onion sets are ready to flower temperatures above 26 7 C 80 F however shifts the sets to the desired bulb forming phase External links edithttps www jic ac uk staff caroline dean vernalization htm Article in New Scientist Retrieved from https en wikipedia org w index php title Vernalization amp oldid 1195484308, wikipedia, wiki, book, books, library,

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