関: vernalization

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出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2018/01/02 23:00:07」(JST)

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Many species of henbane require vernalization before flowering.

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 flower. Vernalization is sometimes used to refer to herbal (non-woody) plants requiring a cold dormancy to produce new shoots and leaves^[1] but this usage is discouraged.^[2]

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 5 and 10 degrees Celsius (40 and 50 degrees Fahrenheit).^{[citation needed]}

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^[4] and winter cereals such as wheat, must go through a prolonged period of cold before flowering occurs.

History of vernalization research

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.^[5] In 1928 a Russian geneticist 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".^[6]

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.^[7] Later however, 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.^[8]

Early research on vernalization focused on plant physiology; the increasing availability of molecular biology has made it possible to unravel its underlying mechanisms.^[8] 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).^[9]

Due to plant flowering requiring the successful co-operation of several metabolic pathways, computer models that incorporate vernalization have also been made.^[10]

In Arabidopsis thaliana

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.^[11] The genes that underlie this difference in plant physiology have been intensively studied.^[8]

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).^[12] Bolting is a robust predictor of flower formation, and hence a good indicator for vernalization research.^[12]

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.^[11]

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]^[13] Winter annual ecotypes of Arabidopsis have an active copy of the gene FRIGIDA (FRI), which promotes FLC expression, thus repression of flowering.^[14] Prolonged exposure to cold (vernalization) induces expression of VERNALIZATION INSENSTIVE3, which interacts with the VERNALIZATION2 polycomb-like complex to reduce FLC expression through chromatin remodeling.^[15] The epigenetic silencing of FLC by chromatin remodeling is also thought to involve the cold-induced expression of antisense FLC COOLAIR^[16]^[17] or COLDAIR transcripts.^[18] Vernalization is registered by the plant by the stable silencing of individual FLC loci.^[19] The removal of silent chromatin marks at FLC during embryogenesis prevents the inheritance of the vernalized state ^[20]

Since vernalization also occurs in flc mutants (lacking FLC), vernalization must also activate a non-FLC pathway.^[21] A day-length mechanism is also important.^[9]

Devernalization

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.^[22]

References

^ 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.
^ 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 & 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.
^ ^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.
^ "New Genetic Model Predicts Plant Flowering in Different Environments" (Press release). Brown University. January 15, 2009. Retrieved March 20, 2016.
^ ^a ^b "Vernalisation response". Plant Biology. Retrieved 2011-01-26. ^{[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 inArabidopsis, 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.
^ http://www.jic.ac.uk/news/2014/10/plants-require-coolair-flower-spring^{[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. doi:10.1126/science.1197349. PMID 21127216.
^ 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.
^ Crevillén, Pedro; Yang, Hongchun; Cui, Xia; Greeff, Christiaan; Trick, Martin; Qiu, Qi; Cao, Xiaofeng; Dean, Caroline (2014). "Epigenetic reprogramming that prevents transgenerational inheritance of the vernalized state". Nature. 515 (7528): 587–90. doi:10.1038/nature13722. PMC 4247276 . PMID 25219852.
^ "Vernalisation pathway". Plant Biology. Retrieved 2011-01-26. ^{[self-published source?]}
^ "Vernalization". Encyclopædia Britannica Online. Retrieved 2011-01-24. vernalization, the artificial exposure of plants (or seeds) to low temperatures in order to stimulate flowering or to enhance seed production. By satisfying the cold requirement of many temperate-zone plants, flowering can be induced to occur earlier than normal or in warm climates lacking the requisite seasonal chilling.

External links

https://www.jic.ac.uk/staff/caroline-dean/vernalization.htm
Article in New Scientist

English Journal

High temperatures limit plant growth but hasten flowering in root chicory (Cichorium intybus) independently of vernalisation.

Mathieu AS1, Lutts S1, Vandoorne B1, Descamps C1, Périlleux C2, Dielen V3, Van Herck JC3, Quinet M4.Author information 1Groupe de Recherche en Physiologie Végétale (GRPV), Earth and Life Institute, Université catholique de Louvain, 5 (bte L7.07.13) Place Croix du Sud, 1348 Louvain-la-Neuve, Belgium.2Université de Liège, Département des Sciences de la Vie PhytoSYSTEMS, Laboratoire de Physiologie Végétale, 27 Boulevard du Rectorat (Bât 22), 4000 Liège, Belgium.3Chicoline - Research and Chicory Breeding, S.A. Warcoing, rue de la Sucrerie 2, B-7740 Warcoing, Belgium.4Groupe de Recherche en Physiologie Végétale (GRPV), Earth and Life Institute, Université catholique de Louvain, 5 (bte L7.07.13) Place Croix du Sud, 1348 Louvain-la-Neuve, Belgium. Electronic address: muriel.quinet@uclouvain.be.AbstractAn increase in mean and extreme summer temperatures is expected as a consequence of climate changes and this might have an impact on plant development in numerous species. Root chicory (Cichorium intybus L.) is a major crop in northern Europe, and it is cultivated as a source of inulin. This polysaccharide is stored in the tap root during the first growing season when the plant grows as a leafy rosette, whereas bolting and flowering occur in the second year after winter vernalisation. The impact of heat stress on plant phenology, water status, photosynthesis-related parameters, and inulin content was studied in the field and under controlled phytotron conditions. In the field, plants of the Crescendo cultivar were cultivated under a closed plastic-panelled greenhouse to investigate heat-stress conditions, while the control plants were shielded with a similar, but open, structure. In the phytotrons, the Crescendo and Fredonia cultivars were exposed to high temperatures (35°C day/28°C night) and compared to control conditions (17°C) over 10 weeks. In the field, heat reduced the root weight, the inulin content of the root and its degree of polymerisation in non-bolting plants. Flowering was observed in 12% of the heat stressed plants during the first growing season in the field. In the phytotron, the heat stress increased the total number of leaves per plant, but reduced the mean leaf area. Photosynthesis efficiency was increased in these plants, whereas osmotic potential was decreased. High temperature was also found to induced flowering of up to 50% of these plants, especially for the Fredonia cultivar. In conclusion, high temperatures induced a reduction in the growth of root chicory, although photosynthesis is not affected. Flowering was also induced, which indicates that high temperatures can partly substitute for the vernalisation requirement for the flowering of root chicory.
Journal of plant physiology.J Plant Physiol.2014 Jan 15;171(2):109-18. doi: 10.1016/j.jplph.2013.09.011. Epub 2013 Nov 20.
An increase in mean and extreme summer temperatures is expected as a consequence of climate changes and this might have an impact on plant development in numerous species. Root chicory (Cichorium intybus L.) is a major crop in northern Europe, and it is cultivated as a source of inulin. This polysac
PMID 24331425

Retroelement insertions at the Medicago FTa1 locus in spring mutants eliminate vernalisation but not long-day requirements for early flowering.

Jaudal M, Yeoh CC, Zhang L, Stockum C, Mysore KS, Ratet P, Putterill J.Author information School of Biological Sciences, The University of Auckland, Private Bag 92019, Auckland Mail Centre, Auckland, 1142, New Zealand.AbstractMolecular-genetic control of the flowering time of temperate-climate plants is best understood in Arabidopsis and the cereals wheat and barley. However, key regulators such as FLC and cereal VRN2 are not found in legumes. Therefore, we used forward genetics to identify flowering time genes in the model legume Medicago truncatula (Medicago) which is induced to flower by vernalisation and long-day photoperiods. A screen of a Tnt1 retroelement tagging population yielded two mutants, spring2 and spring3, with a dominant early flowering phenotype. These mutants overexpress the floral activator FTa1 and two candidate downstream flowering genes SOC1a and FULb, similar to the spring1 somaclonal variant that we identified previously. We demonstrate here that an increase in the expression of FTa1, SOC1a and FULb and early flowering does not occur in all conditions in the spring mutants. It depends on long-day photoperiods but not on vernalisation. Isolation of flanking sequence tags and linkage analysis identified retroelement insertions at FTa1 that co-segregated with the early flowering phenotype in all three spring mutants. These were Tnt1 insertions in the FTa1 third intron (spring3) or the 3' intergenic region (spring2) and an endogenous MERE1-4 retroelement in the 3' intergenic region in spring1. Thus the spring mutants form an allelic series of gain-of-function mutations in FTa1 which confer a spring growth habit. The spring retroelement insertions at FTa1 separate long-day input from vernalisation input into FTa1 regulation, but this is not due to large-scale changes in FTa1 DNA methylation or transcript processing in the mutants.
The Plant journal : for cell and molecular biology.Plant J.2013 Nov;76(4):580-91. doi: 10.1111/tpj.12315. Epub 2013 Oct 5.
Molecular-genetic control of the flowering time of temperate-climate plants is best understood in Arabidopsis and the cereals wheat and barley. However, key regulators such as FLC and cereal VRN2 are not found in legumes. Therefore, we used forward genetics to identify flowering time genes in the mo
PMID 23964816

Variation at the vernalisation genes Vrn-H1 and Vrn-H2 determines growth and yield stability in barley (Hordeum vulgare) grown under dryland conditions in Syria.

Rollins JA, Drosse B, Mulki MA, Grando S, Baum M, Singh M, Ceccarelli S, von Korff M.Author information Max Planck Institute for Plant Breeding Research, Carl-von-Linné-Weg 10, 50829, Cologne, Germany.AbstractKEY MESSAGE: Spring growth in barley controlled by natural variation at Vrn-H1 and Vrn-H2 improved yield stability in marginal Syrian environments. The objective of the present study was to identify QTL influencing agronomic performance in rain-fed Mediterranean environments in a recombinant inbred line (RIL) population, ARKE derived from the Syrian barley landrace, Arta and the Australian feed cultivar, Keel. The population was field tested for agronomic performance at two locations in Syria for 4 years with two sowing dates, in autumn and winter. Genotypic variability in yield of the RIL population was mainly affected by year-to-year variation presumably caused by inter-annual differences in rainfall distribution. The spring growth habit and early flowering inherited from the Australian cultivar Keel increased plant height and biomass and improved yield stability in Syrian environments. QTL for yield and biomass coincided with the map location of flowering time genes, in particular the vernalisation genes Vrn-H1 and Vrn-H2. In marginal environments with terminal drought, the Vrn-H1 allele inherited from Keel improved final biomass and yield. Under changing climate conditions, such as shorter winters, reduced rainfall, and early summer drought, spring barley might thus outperform the traditional vernalisation-sensitive Syrian landraces. We present the ARKE population as a valuable genetic resource to further elucidate the genetics of drought adaptation of barley in the field.
TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik.Theor Appl Genet.2013 Nov;126(11):2803-24. doi: 10.1007/s00122-013-2173-y. Epub 2013 Aug 6.
KEY MESSAGE: Spring growth in barley controlled by natural variation at Vrn-H1 and Vrn-H2 improved yield stability in marginal Syrian environments. The objective of the present study was to identify QTL influencing agronomic performance in rain-fed Mediterranean environments in a recombinant inbred
PMID 23918065

Japanese Journal

Synchronisation of Arabidopsis flowering time and whole-plant senescence in seasonal environments

Miryeganeh Matin,Yamaguchi Masaki,Kudoh Hiroshi
Scientific Reports (8), 2018-07-06
… The role of vernalisation in realising flowering synchrony between plants of different ages under natural conditions was demonstrated by synchronisation and de-synchronisation of flowering initiation in vernalisation-sensitive and less-vernalisation-sensitive accessions, respectively. …
NAID 120006502584

海浜植物群落の保全と復元について：系統地理学と集団遺伝学からの提言

瀬戸口浩彰
景観生態学 22(2), 43-51, 2017
<p>海浜植物には種子散布が海流によって長距離に及ぶものが多く，また匍匐茎が拡がって多くのラメットを形成するものも多く存在する．そのために，海浜植物は系統地理構造を作りにくく，遺伝的な多様性が低いというイメージがある．本稿では，DNAデータを用いた系統地理学と集団遺伝学のいくつかの知見をもとにして，海浜植物群落を保全や復元するにあたっての注意点を提起した．ハマヒルガオとハマエンドウでは …
NAID 130007667127

Lineage isolation in the face of active gene flow in the coastal plant wild radish is reinforced by differentiated vernalisation responses

Han Qingxiang,Higashi Hiroyuki,Mitsui Yuki,Setoguchi Hiroaki
BMC Evolutionary Biology (16), 2016-04-16
… In addition, we investigated the flowering times of individuals in different wild radish lineages, with and without cold exposure, to assess their respective vernalisation responses. … Our greenhouse experiments indicated that cold exposure was not required for flowering in the southern group, but could advance the date of flowering, suggesting that vernalisation would be facultative in the southern group. …
NAID 120005947144

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関: vernalisation

[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.

[chouard-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.

[sung-3] 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.

[4] 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.

[5] 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.

[6] Poltronieri, Palmiro; Hong, Yiguo (2015). Applied Plant Genomics and Biotechnology. Cambridge (UK): Woodhead Publishing. p. 121. ISBN 978-0-08-100068-7.

[lysenkosci-7] Roll-Hansen, Nils (1985). "A new perspective on Lysenko?". Annals of Science. Taylor & Francis. 42 (3): 261–278. doi:10.1080/00033798500200201. PMID 11620694.

[amasino-8] 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.

[trevaskis-9] 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.

[10] "New Genetic Model Predicts Plant Flowering in Different Environments" (Press release). Brown University. January 15, 2009. Retrieved March 20, 2016.

[pbvr-11] "Vernalisation response". Plant Biology. Retrieved 2011-01-26. ^{[self-published source?]}

[pouteau-12] 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.

[13] 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.

[14] 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 inArabidopsis, by Recruiting Chromatin Modification Factors". The Plant Cell. 23 (1): 289–303. doi:10.1105/tpc.110.075911. PMC 3051252 . PMID 21282526.

[Sung_and_Amasino-15] 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.

[16] ttp://www.jic.ac.uk/news/2014/10/plants-require-coolair-flower-spring^{[full citation needed]}

[Csorba-17] 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.

[pmid21127216-18] Heo, J. B.; Sung, S. (2011). "Vernalization-Mediated Epigenetic Silencing by a Long Intronic Noncoding RNA". Science. 331 (6013): 76–9. doi:10.1126/science.1197349. PMID 21127216.

[Angel-19] 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.

[Crevillen-20] Crevillén, Pedro; Yang, Hongchun; Cui, Xia; Greeff, Christiaan; Trick, Martin; Qiu, Qi; Cao, Xiaofeng; Dean, Caroline (2014). "Epigenetic reprogramming that prevents transgenerational inheritance of the vernalized state". Nature. 515 (7528): 587–90. doi:10.1038/nature13722. PMC 4247276 . PMID 25219852.

[pbvp-21] "Vernalisation pathway". Plant Biology. Retrieved 2011-01-26. ^{[self-published source?]}

[22] "Vernalization". Encyclopædia Britannica Online. Retrieved 2011-01-24. vernalization, the artificial exposure of plants (or seeds) to low temperatures in order to stimulate flowering or to enhance seed production. By satisfying the cold requirement of many temperate-zone plants, flowering can be induced to occur earlier than normal or in warm climates lacking the requisite seasonal chilling.

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Concepts	Heredity Heritability Hybridization Inheritance of acquired characteristics Lamarckism Mendelian inheritance Vernalization
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