1932

Abstract

Many plants use information about changing day length (photoperiod) to align their flowering time with seasonal changes to increase reproductive success. A mechanism for photoperiodic time measurement is present in leaves, and the day-length-specific induction of the () gene, which encodes florigen, is a major final output of the pathway. Here, we summarize the current understanding of the molecular mechanisms by which photoperiodic information is perceived in order to trigger expression in as well as in the primary cereals wheat, barley, and rice. In these plants, the differences in photoperiod are measured by interactions between circadian-clock-regulated components, such as CONSTANS (CO), and light signaling. The interactions happen under certain day-length conditions, as previously predicted by the external coincidence model. In these plants, the coincidence mechanisms are governed by multilayered regulation with numerous conserved as well as unique regulatory components, highlighting the breadth of photoperiodic regulation across plant species.

Loading

Article metrics loading...

/content/journals/10.1146/annurev-arplant-043014-115555
2015-04-29
2024-04-29
Loading full text...

Full text loading...

/deliver/fulltext/arplant/66/1/annurev-arplant-043014-115555.html?itemId=/content/journals/10.1146/annurev-arplant-043014-115555&mimeType=html&fmt=ahah

Literature Cited

  1. Alon U. 1.  2007. Network motifs: theory and experimental approaches. Nat. Rev. Genet. 8:450–61 [Google Scholar]
  2. An H, Roussot C, Suárez-López P, Corbesier L, Vincent C. 2.  et al. 2004. CONSTANS acts in the phloem to regulate a systemic signal that induces photoperiodic flowering of Arabidopsis. Development 131:3615–26 [Google Scholar]
  3. Andrés F, Coupland G. 3.  2012. The genetic basis of flowering responses to seasonal cues. Nat. Rev. Genet. 13:627–39 [Google Scholar]
  4. Beales J, Turner A, Griffiths S, Snape JW, Laurie DA. 4.  2007. A Pseudo-Response Regulator is misexpressed in the photoperiod insensitive Ppd-D1a mutant of wheat (Triticum aestivum L.). Theor. Appl. Genet. 115:721–33 [Google Scholar]
  5. Brooking IR, Jamieson PD. 5.  2002. Temperature and photoperiod response of vernalization in near-isogenic lines of wheat. Field Crops Res. 79:21–38 [Google Scholar]
  6. Brown HE, Jamieson PD, Brooking IR, Moot DJ, Huth NI. 6.  2013. Integration of molecular and physiological models to explain time of anthesis in wheat. Ann. Bot. 112:1683–703 [Google Scholar]
  7. Budhiraja R, Hermkes R, Muller S, Schmidt J, Colby T. 7.  et al. 2009. Substrates related to chromatin and to RNA-dependent processes are modified by Arabidopsis SUMO isoforms that differ in a conserved residue with influence on desumoylation. Plant Physiol. 149:1529–40 [Google Scholar]
  8. Bünning E. 8.  1960. Opening address: biological clocks. Cold Spring Harb. Symp. Quant. Biol. 25:1–9 [Google Scholar]
  9. Campoli C, Shtaya M, Davis SJ, von Korff M. 9.  2012. Expression conservation within the circadian clock of a monocot: natural variation at barley Ppd-H1 affects circadian expression of flowering time genes, but not clock orthologs. BMC Plant Biol. 12:97 [Google Scholar]
  10. Cao S, Kumimoto RW, Gnesutta N, Calogero AM, Mantovani R, Holt BF III. 10.  2014. A distal CCAAT/NUCLEAR FACTOR Y complex promotes chromatin looping at the FLOWERING LOCUS T promoter and regulates the timing of flowering in Arabidopsis. Plant Cell 26:1009–17 [Google Scholar]
  11. Castillejo C, Pelaz S. 11.  2008. The balance between CONSTANS and TEMPRANILLO activities determines FT expression to trigger flowering. Curr. Biol. 18:1338–43 [Google Scholar]
  12. Chen A, Dubcovsky J. 12.  2012. Wheat TILLING mutants show that the vernalization gene VRN1 down-regulates the flowering repressor VRN2 in leaves but is not essential for flowering. PLOS Genet. 8:e1003134 [Google Scholar]
  13. Chen A, Li C, Hu W, Lau MY, Lin H. 13.  et al. 2014. PHYTOCHROME C plays a major role in the acceleration of wheat flowering under long-day photoperiod. PNAS 111:10037–44 [Google Scholar]
  14. Chen J, Li X, Cheng C, Wang Y, Qin M. 14.  et al. 2014. Characterization of epistatic interaction of QTLs LH8 and EH3 controlling heading date in rice. Sci. Rep. 4:4263 [Google Scholar]
  15. Choi SC, Lee S, Kim SR, Lee YS, Liu C. 15.  et al. 2014. Trithorax group protein Oryza sativa Trithorax1 controls flowering time in rice via interaction with Early heading date3. Plant Physiol. 164:1326–37 [Google Scholar]
  16. Clack T, Shokry A, Moffet M, Liu P, Faul M, Sharrock RA. 16.  2009. Obligate heterodimerization of Arabidopsis phytochromes C and E and interaction with the PIF3 basic helix-loop-helix transcription factor. Plant Cell 21:786–99 [Google Scholar]
  17. Cumming BG, Hendricks SB, Borthwick HA. 17.  1965. Rhythmic flowering responses and phytochrome changes in a selection of Chenopodium rubrum. Can. J. Bot. 43:825–53 [Google Scholar]
  18. David KM, Armbruster U, Tama N, Putterill J. 18.  2006. Arabidopsis GIGANTEA protein is post-transcriptionally regulated by light and dark. FEBS Lett. 580:1193–97 [Google Scholar]
  19. Díaz A, Zikhali M, Turner AS, Isaac P, Laurie DA. 19.  2012. Copy number variation affecting the Photoperiod-B1 and Vernalization-A1 genes is associated with altered flowering time in wheat (Triticum aestivum). PLOS ONE 7:e33234 [Google Scholar]
  20. Distelfeld A, Dubcovsky J. 20.  2010. Characterization of the maintained vegetative phase deletions from diploid wheat and their effect on VRN2 and FT transcript levels. Mol. Genet. Genomics 283:223–32 [Google Scholar]
  21. Distelfeld A, Li C, Dubcovsky J. 21.  2009. Regulation of flowering in temperate cereals. Curr. Opin. Plant Biol. 12:178–84 [Google Scholar]
  22. Doi K, Izawa T, Fuse T, Yamanouchi U, Kubo T. 22.  et al. 2004. Ehd1, a B-type response regulator in rice, confers short-day promotion of flowering and controls FT-like gene expression independently of Hd1. Genes Dev. 18:926–36 [Google Scholar]
  23. Dubcovsky J, Loukoianov A, Fu DL, Valarik M, Sanchez A, Yan LL. 23.  2006. Effect of photoperiod on the regulation of wheat vernalization genes VRN1 and VRN2. Plant Mol. Biol. 60:469–80 [Google Scholar]
  24. Elrouby N, Bonequi MV, Porri A, Coupland G. 24.  2013. Identification of Arabidopsis SUMO-interacting proteins that regulate chromatin activity and developmental transitions. PNAS 110:19956–61 [Google Scholar]
  25. Endo M, Tanigawa Y, Murakami T, Araki T, Nagatani A. 25.  2013. PHYTOCHROME-DEPENDENT LATE-FLOWERING accelerates flowering through physical interactions with phytochrome B and CONSTANS. PNAS 110:18017–22 [Google Scholar]
  26. Fornara F, Panigrahi KC, Gissot L, Sauerbrunn N, Rühl M. 26.  et al. 2009. Arabidopsis DOF transcription factors act redundantly to reduce CONSTANS expression and are essential for a photoperiodic flowering response. Dev. Cell 17:75–86 [Google Scholar]
  27. Fowler S, Lee K, Onouchi H, Samach A, Richardson K. 27.  et al. 1999. GIGANTEA: a circadian clock-controlled gene that regulates photoperiodic flowering in Arabidopsis and encodes a protein with several possible membrane-spanning domains. EMBO J. 18:4679–88 [Google Scholar]
  28. Gao H, Zheng X-M, Fei G, Chen J, Jin M. 28.  et al. 2013. Ehd4 encodes a novel and Oryza-genus-specific regulator of photoperiodic flowering in rice. PLOS Genet. 9:e1003281 [Google Scholar]
  29. Garner WW, Allard HA. 29.  1920. Effect of the relative length of day and night and other factors of the environment on growth and reproduction in plants. J. Agric. 18:553–606 [Google Scholar]
  30. Gendron JM, Pruneda-Paz JL, Doherty CJ, Gross AM, Kang SE, Kay SA. 30.  2012. Arabidopsis circadian clock protein, TOC1, is a DNA-binding transcription factor. PNAS 109:3167–72 [Google Scholar]
  31. Giakountis A, Coupland G. 31.  2008. Phloem transport of flowering signals. Curr. Opin. Plant Biol. 11:687–94 [Google Scholar]
  32. Gomez D, Vanzetti L, Helguera M, Lombardo L, Fraschina J, Miralles DJ. 32.  2014. Effect of Vrn-1, Ppd-1 genes and earliness per se on heading time in Argentinean bread wheat cultivars. Field Crops Res. 158:73–81 [Google Scholar]
  33. Hayama R, Agashe B, Luley E, King R, Coupland G. 33.  2007. A circadian rhythm set by dusk determines the expression of FT homologs and the short-day photoperiodic flowering response in Pharbitis. Plant Cell 19:2988–3000 [Google Scholar]
  34. Hayama R, Yokoi S, Tamaki S, Yano M, Shimamoto K. 34.  2003. Adaptation of photoperiodic control pathways produces short-day flowering in rice. Nature 422:719–22 [Google Scholar]
  35. He Y. 35.  2012. Chromatin regulation of flowering. Trends Plant Sci. 17:556–62 [Google Scholar]
  36. Hori K, Ogiso-Tanaka E, Matsubara K, Yamanouchi U, Ebana K, Yano M. 36.  2013. Hd16, a gene for casein kinase I, is involved in the control of rice flowering time by modulating the day-length response. Plant J. 76:36–46 [Google Scholar]
  37. Huang W, Pérez-García P, Pokhilko A, Millar AJ, Antoshechkin I. 37.  et al. 2012. Mapping the core of the Arabidopsis circadian clock defines the network structure of the oscillator. Science 336:75–79 [Google Scholar]
  38. Imaizumi T. 38.  2010. Arabidopsis circadian clock and photoperiodism: time to think about location. Curr. Opin. Plant Biol. 13:83–89 [Google Scholar]
  39. Imaizumi T, Schultz TF, Harmon FG, Ho LA, Kay SA. 39.  2005. FKF1 F-box protein mediates cyclic degradation of a repressor of CONSTANS in Arabidopsis. Science 309:293–97 [Google Scholar]
  40. Imaizumi T, Tran HG, Swartz TE, Briggs WR, Kay SA. 40.  2003. FKF1 is essential for photoperiodic-specific light signalling in Arabidopsis. Nature 426:302–6 [Google Scholar]
  41. Ishikawa R, Aoki M, Kurotani K, Yokoi S, Shinomura T. 41.  et al. 2011. Phytochrome B regulates Heading date 1 (Hd1)-mediated expression of rice florigen Hd3a and critical day length in rice. Mol. Genet. Genomics 285:461–70 [Google Scholar]
  42. Ito S, Niwa Y, Nakamichi N, Kawamura H, Yamashino T, Mizuno T. 42.  2008. Insight into missing genetic links between two evening-expressed pseudo-response regulator genes TOC1 and PRR5 in the circadian clock-controlled circuitry in Arabidopsis thaliana. Plant Cell Physiol. 49:201–13 [Google Scholar]
  43. Ito S, Song YH, Josephson-Day AR, Miller RJ, Breton G. 43.  et al. 2012. FLOWERING BHLH transcriptional activators control expression of the photoperiodic flowering regulator CONSTANS in Arabidopsis. PNAS 109:3582–87 [Google Scholar]
  44. Itoh H, Nonoue Y, Yano M, Izawa T. 44.  2010. A pair of floral regulators sets critical day length for Hd3a florigen expression in rice. Nat. Genet. 42:635–38 [Google Scholar]
  45. Izawa T. 45.  2007. Adaptation of flowering-time by natural and artificial selection in Arabidopsis and rice. J. Exp. Bot. 58:3091–97 [Google Scholar]
  46. Izawa T, Oikawa T, Sugiyama N, Tanisaka T, Yano M, Shimamoto K. 46.  2002. Phytochrome mediates the external light signal to repress FT orthologs in photoperiodic flowering of rice. Genes Dev. 16:2006–20 [Google Scholar]
  47. Jang S, Marchal V, Panigrahi KC, Wenkel S, Soppe W. 47.  et al. 2008. Arabidopsis COP1 shapes the temporal pattern of CO accumulation conferring a photoperiodic flowering response. EMBO J. 27:1277–88 [Google Scholar]
  48. Jeong J-H, Song H-R, Ko J-H, Jeong Y-M, Kwon YE. 48.  et al. 2009. Repression of FLOWERING LOCUS T chromatin by functionally redundant histone H3 lysine 4 demethylases in Arabidopsis. PLOS ONE 4:e8033 [Google Scholar]
  49. Jung JH, Seo YH, Seo PJ, Reyes JL, Yun J. 49.  et al. 2007. The GIGANTEA-regulated microRNA172 mediates photoperiodic flowering independent of CONSTANS in Arabidopsis. Plant Cell 19:2736–48 [Google Scholar]
  50. Kasperbauer MJ, Borthwick HA, Hendricks SB. 50.  1963. Inhibition of flowering of Chenopodium rubrum by prolonged far-red radiation. Bot. Gaz. 124:444–51 [Google Scholar]
  51. Kim WY, Fujiwara S, Suh SS, Kim J, Kim Y. 51.  et al. 2007. ZEITLUPE is a circadian photoreceptor stabilized by GIGANTEA in blue light. Nature 449:356–60 [Google Scholar]
  52. Kitagawa S, Shimada S, Murai K. 52.  2012. Effect of Ppd-1 on the expression of flowering-time genes in vegetative and reproductive growth stages of wheat. Genes Genet. Syst. 87:161–68 [Google Scholar]
  53. Kobayashi Y, Kaya H, Goto K, Iwabuchi M, Araki T. 53.  1999. A pair of related genes with antagonistic roles in mediating flowering signals. Science 286:1960–62 [Google Scholar]
  54. Komiya R, Ikegami A, Tamaki S, Yokoi S, Shimamoto K. 54.  2008. Hd3a and RFT1 are essential for flowering in rice. Development 135:767–74 [Google Scholar]
  55. Komiya R, Yokoi S, Shimamoto K. 55.  2009. A gene network for long-day flowering activates RFT1 encoding a mobile flowering signal in rice. Development 136:3443–50 [Google Scholar]
  56. Koo BH, Yoo SC, Park JW, Kwon CT, Lee BD. 56.  et al. 2013. Natural variation in OsPRR37 regulates heading date and contributes to rice cultivation at a wide range of latitudes. Mol. Plant 6:1877–88 [Google Scholar]
  57. Kubota A, Kita S, Ishizaki K, Nishihama R, Yamato KT, Kohchi T. 57.  2014. Co-option of a photoperiodic growth-phase transition system during land plant evolution. Nat. Commun. 5:3668 [Google Scholar]
  58. Kumimoto RW, Adam L, Hymus GJ, Repetti PP, Reuber TL. 58.  et al. 2008. The Nuclear Factor Y subunits NF-YB2 and NF-YB3 play additive roles in the promotion of flowering by inductive long-day photoperiods in Arabidopsis. Planta 228:709–23 [Google Scholar]
  59. Kumimoto RW, Zhang Y, Siefers N, Holt BF. 59.  2010. NF-YC3, NF-YC4 and NF-YC9 are required for CONSTANS-mediated, photoperiod-dependent flowering in Arabidopsis thaliana. Plant J. 63:379–91 [Google Scholar]
  60. Lanning SP, Hucl P, Pumphrey M, Carter AH, Lamb PF. 60.  et al. 2012. Agronomic performance of spring wheat as related to planting date and photoperiod response. Crop Sci. 52:1633–39 [Google Scholar]
  61. Laubinger S, Marchal V, Le Gourrierec J, Wenkel S, Adrian J. 61.  et al. 2006. Arabidopsis SPA proteins regulate photoperiodic flowering and interact with the floral inducer CONSTANS to regulate its stability. Development 133:3213–22 [Google Scholar]
  62. Lazaro A, Valverde F, Piñeiro M, Jarillo JA. 62.  2012. The Arabidopsis E3 ubiquitin ligase HOS1 negatively regulates CONSTANS abundance in the photoperiodic control of flowering. Plant Cell 24:982–99 [Google Scholar]
  63. Lee YS, Jeong DH, Lee DY, Yi J, Ryu CH. 63.  et al. 2010. OsCOL4 is a constitutive flowering repressor upstream of Ehd1 and downstream of OsphyB. Plant J. 63:18–30 [Google Scholar]
  64. Li CX, Dubcovsky J. 64.  2008. Wheat FT protein regulates VRN1 transcription through interactions with FDL2. Plant J. 55:543–54 [Google Scholar]
  65. Li D, Yang C, Li X, Gan Q, Zhao X, Zhu L. 65.  2009. Functional characterization of rice OsDof12. Planta 229:1159–69 [Google Scholar]
  66. Liu H, Wang Q, Liu Y, Zhao X, Imaizumi T. 66.  et al. 2013. Arabidopsis CRY2 and ZTL mediate blue-light regulation of the transcription factor CIB1 by distinct mechanisms. PNAS 110:17582–87 [Google Scholar]
  67. Liu H, Yu X, Li K, Klejnot J, Yang H. 67.  et al. 2008. Photoexcited CRY2 interacts with CIB1 to regulate transcription and floral initiation in Arabidopsis. Science 322:1535–39 [Google Scholar]
  68. Liu L, Wang E, Zhu Y, Tang L, Cao W. 68.  2013. Quantifying three-decade changes of single rice cultivars in China using crop modeling. Field Crops Res. 149:84–94 [Google Scholar]
  69. Liu L, Zhu Y, Shen L, Yu H. 69.  2013. Emerging insights into florigen transport. Curr. Opin. Plant Biol. 16:607–13 [Google Scholar]
  70. Liu LJ, Zhang YC, Li QH, Sang Y, Mao J. 70.  et al. 2008. COP1-mediated ubiquitination of CONSTANS is implicated in cryptochrome regulation of flowering in Arabidopsis. Plant Cell 20:292–306 [Google Scholar]
  71. Liu Y, Li X, Li K, Liu H, Lin C. 71.  2013. Multiple bHLH proteins form heterodimers to mediate CRY2-dependent regulation of flowering-time in Arabidopsis. PLOS Genet. 9:e1003861 [Google Scholar]
  72. Mathieu J, Yant LJ, Mürdter F, Küttner F, Schmid M. 72.  2009. Repression of flowering by the miR172 target SMZ. PLOS Biol. 7:e1000148 [Google Scholar]
  73. Matsoukas IG, Massiah AJ, Thomas B. 73.  2013. Starch metabolism and antiflorigenic signals modulate the juvenile-to-adult phase transition in Arabidopsis. Plant Cell Environ. 36:1802–11 [Google Scholar]
  74. Matsubara K, Ogiso-Tanaka E, Hori K, Ebana K, Ando T, Yano M. 74.  2012. Natural variation in Hd17, a homolog of Arabidopsis ELF3 that is involved in rice photoperiodic flowering. Plant Cell Physiol. 53:709–16 [Google Scholar]
  75. Matsubara K, Yamanouchi U, Nonoue Y, Sugimoto K, Wang ZX. 75.  et al. 2011. Ehd3, encoding a plant homeodomain finger-containing protein, is a critical promoter of rice flowering. Plant J. 66:603–12 [Google Scholar]
  76. Matsubara K, Yamanouchi U, Wang ZX, Minobe Y, Izawa T, Yano M. 76.  2008. Ehd2, a rice ortholog of the maize INDETERMINATE1 gene, promotes flowering by up-regulating Ehd1. Plant Physiol. 148:1425–35 [Google Scholar]
  77. Mizuno N, Nitta M, Sato K, Nasuda S. 77.  2012. A wheat homologue of PHYTOCLOCK 1 is a candidate gene conferring the early heading phenotype to einkorn wheat. Genes Genet. Syst. 87:357–67 [Google Scholar]
  78. Moglich A, Yang X, Ayers RA, Moffat K. 78.  2010. Structure and function of plant photoreceptors. Annu. Rev. Plant Biol. 61:21–47 [Google Scholar]
  79. Murakami M, Tago Y, Yamashino T, Mizuno T. 79.  2007. Comparative overviews of clock-associated genes of Arabidopsis thaliana and Oryza sativa. Plant Cell Physiol. 48:110–21 [Google Scholar]
  80. Nakamichi N, Kiba T, Henriques R, Mizuno T, Chua NH, Sakakibara H. 80.  2010. PSEUDO-RESPONSE REGULATORS 9, 7, and 5 are transcriptional repressors in the Arabidopsis circadian clock. Plant Cell 22:594–605 [Google Scholar]
  81. Nakamichi N, Kiba T, Kamioka M, Suzuki T, Yamashino T. 81.  et al. 2012. Transcriptional repressor PRR5 directly regulates clock-output pathways. PNAS 109:17123–28 [Google Scholar]
  82. Nakamichi N, Kita M, Niimura K, Ito S, Yamashino T. 82.  et al. 2007. Arabidopsis clock-associated pseudo-response regulators PRR9, PRR7 and PRR5 coordinately and positively regulate flowering time through the canonical CONSTANS-dependent photoperiodic pathway. Plant Cell Physiol. 48:822–32 [Google Scholar]
  83. Nishida H, Ishihara D, Ishii M, Kaneko T, Kawahigashi H. 83.  et al. 2013. Phytochrome C is a key factor controlling long-day flowering in barley (Hordeum vulgare L.). Plant Physiol. 163:804–14 [Google Scholar]
  84. Nishida H, Yoshida T, Kawakami K, Fujita M, Long B. 84.  et al. 2013. Structural variation in the 5′ upstream region of photoperiod-insensitive alleles Ppd-A1a and Ppd-B1a identified in hexaploid wheat (Triticum aestivum L.), and their effect on heading time. Mol. Breed. 31:27–37 [Google Scholar]
  85. Ogiso E, Takahashi Y, Sasaki T, Yano M, Izawa T. 85.  2010. The role of casein kinase II in flowering time regulation has diversified during evolution. Plant Physiol. 152:808–20 [Google Scholar]
  86. Osugi A, Itoh H, Ikeda-Kawakatsu K, Takano M, Izawa T. 86.  2011. Molecular dissection of the roles of phytochrome in photoperiodic flowering in rice. Plant Physiol. 157:1128–37 [Google Scholar]
  87. Peng LT, Shi ZY, Li L, Shen GZ, Zhang JL. 87.  2007. Ectopic expression of OsLFL1 in rice represses Ehd1 by binding on its promoter. Biochem. Biophys. Res. Commun. 360:251–56 [Google Scholar]
  88. Peng LT, Shi ZY, Li L, Shen GZ, Zhang JL. 88.  2008. Overexpression of transcription factor OsLFL1 delays flowering time in Oryza sativa. J. Plant Physiol. 165:876–85 [Google Scholar]
  89. Pin PA, Nilsson O. 89.  2012. The multifaceted roles of FLOWERING LOCUS T in plant development. Plant Cell Environ. 35:1742–55 [Google Scholar]
  90. Pittendrigh CS. 90.  1966. The circadian oscillation in Drosophila pseudoobscura pupae: a model for the photoperiodic clock. Z. Pflanzenphysiol. 54:275–307 [Google Scholar]
  91. Pittendrigh CS. 91.  1993. Temporal organization: reflections of a Darwinian clock-watcher. Annu. Rev. Physiol. 55:16–54 [Google Scholar]
  92. Pittendrigh CS, Minis DH. 92.  1964. The entrainment of circadian oscillations by light and their role as photoperiodic clocks. Am. Nat. 98:261–94 [Google Scholar]
  93. Putterill J, Robson F, Lee K, Simon R, Coupland G. 93.  1995. The CONSTANS gene of Arabidopsis promotes flowering and encodes a protein showing similarities to zinc finger transcription factors. Cell 80:847–57 [Google Scholar]
  94. Ranjan A, Dickopf S, Ullrich KK, Rensing SA, Hoecker U. 94.  2014. Functional analysis of COP1 and SPA orthologs from Physcomitrella and rice during photomorphogenesis of transgenic Arabidopsis reveals distinct evolutionary conservation. BMC Plant Biol. 14:178 [Google Scholar]
  95. Robson F, Costa MM, Hepworth SR, Vizir I, Piñeiro M. 95.  et al. 2001. Functional importance of conserved domains in the flowering-time gene CONSTANS demonstrated by analysis of mutant alleles and transgenic plants. Plant J. 28:619–31 [Google Scholar]
  96. Rosas U, Mei Y, Xie Q, Banta JA, Zhou RW. 96.  et al. 2014. Variation in Arabidopsis flowering time associated with cis-regulatory variation in CONSTANS. Nat. Commun. 5:3651 [Google Scholar]
  97. Ryu C-H, Lee S, Cho L-H, Kim SL, Lee Y-S. 97.  et al. 2009. OsMADS50 and OsMADS56 function antagonistically in regulating long day (LD)-dependent flowering in rice. Plant Cell Environ. 32:1412–27 [Google Scholar]
  98. Sacks WJ, Kucharik CJ. 98.  2011. Crop management and phenology trends in the U.S. Corn Belt: impacts on yields, evapotranspiration and energy balance. Agric. For. Meteorol. 151:882–94 [Google Scholar]
  99. Saijo Y, Sullivan JA, Wang H, Yang J, Shen Y. 99.  et al. 2003. The COP1-SPA1 interaction defines a critical step in phytochrome A-mediated regulation of HY5 activity. Genes Dev. 17:2642–47 [Google Scholar]
  100. Saito H, Ogiso-Tanaka E, Okumoto Y, Yoshitake Y, Izumi H. 100.  et al. 2012. Ef7 encodes an ELF3-like protein and promotes rice flowering by negatively regulating the floral repressor gene Ghd7 under both short- and long-day conditions. Plant Cell Physiol. 53:717–28 [Google Scholar]
  101. Samach A, Onouchi H, Gold SE, Ditta GS, Schwarz-Sommer Z. 101.  et al. 2000. Distinct roles of CONSTANS target genes in reproductive development of Arabidopsis. Science 288:1613–16 [Google Scholar]
  102. Sawa M, Kay SA. 102.  2011. GIGANTEA directly activates Flowering Locus T in Arabidopsis thaliana. PNAS 108:11698–703 [Google Scholar]
  103. Sawa M, Nusinow DA, Kay SA, Imaizumi T. 103.  2007. FKF1 and GIGANTEA complex formation is required for day-length measurement in Arabidopsis. Science 318:261–65 [Google Scholar]
  104. Schaffer R, Ramsay N, Samach A, Corden S, Putterill J. 104.  et al. 1998. The late elongated hypocotyl mutation of Arabidopsis disrupts circadian rhythms and the photoperiodic control of flowering. Cell 93:1219–29 [Google Scholar]
  105. Shaw LM, Turner AS, Herry L, Griffiths S, Laurie DA. 105.  2013. Mutant alleles of Photoperiod-1 in wheat (Triticum aestivum L.) that confer a late flowering phenotype in long days. PLOS ONE 8:e79459 [Google Scholar]
  106. Shaw LM, Turner AS, Laurie DA. 106.  2012. The impact of photoperiod insensitive Ppd-1a mutations on the photoperiod pathway across the three genomes of hexaploid wheat (Triticum aestivum). Plant J. 71:71–84 [Google Scholar]
  107. Shimada S, Ogawa T, Kitagawa S, Suzuki T, Ikari C. 107.  et al. 2009. A genetic network of flowering-time genes in wheat leaves, in which an APETALA1/FRUITFULL-like gene, VRN1, is upstream of FLOWERING LOCUS T. Plant J. 58:668–81 [Google Scholar]
  108. Shrestha R, Gómez-Ariza J, Brambilla V, Fornara F. 108.  2014. Molecular control of seasonal flowering in rice, arabidopsis and temperate cereals. Ann. Bot. 114:1445–58 [Google Scholar]
  109. Slafer G, Rawson H. 109.  1994. Sensitivity of wheat phasic development to major environmental factors: a re-examination of some assumptions made by physiologists and modellers. Funct. Plant Biol. 21:393–426 [Google Scholar]
  110. Song YH, Ito S, Imaizumi T. 110.  2010. Similarities in the circadian clock and photoperiodism in plants. Curr. Opin. Plant Biol. 13:594–603 [Google Scholar]
  111. Song YH, Ito S, Imaizumi T. 111.  2013. Flowering time regulation: photoperiod- and temperature-sensing in leaves. Trends Plant Sci. 18:575–83 [Google Scholar]
  112. Song YH, Smith RW, To BJ, Millar AJ, Imaizumi T. 112.  2012. FKF1 conveys timing information for CONSTANS stabilization in photoperiodic flowering. Science 336:1045–49 [Google Scholar]
  113. Suárez-López P, Wheatley K, Robson F, Onouchi H, Valverde F, Coupland G. 113.  2001. CONSTANS mediates between the circadian clock and the control of flowering in Arabidopsis. Nature 410:1116–20 [Google Scholar]
  114. Sun C, Fang J, Zhao T, Xu B, Zhang F. 114.  et al. 2012. The histone methyltransferase SDG724 mediates H3K36me2/3 deposition at MADS50 and RFT1 and promotes flowering in rice. Plant Cell 24:3235–47 [Google Scholar]
  115. Takada S, Goto K. 115.  2003. TERMINAL FLOWER2, an Arabidopsis homolog of HETERO-CHROMATIN PROTEIN1, counteracts the activation of FLOWERING LOCUS T by CONSTANS in the vascular tissues of leaves to regulate flowering time. Plant Cell 15:2856–65 [Google Scholar]
  116. Takahashi Y, Shomura A, Sasaki T, Yano M. 116.  2001. Hd6, a rice quantitative trait locus involved in photoperiod sensitivity, encodes the alpha subunit of protein kinase CK2. PNAS 98:7922–27 [Google Scholar]
  117. Tamaki S, Matsuo S, Wong HL, Yokoi S, Shimamoto K. 117.  2007. Hd3a protein is a mobile flowering signal in rice. Science 316:1033–36 [Google Scholar]
  118. Tan S-T, Dai C, Liu H-T, Xue H-W. 118.  2013. Arabidopsis casein kinase1 proteins CK1.3 and CK1.4 phosphorylate cryptochrome2 to regulate blue light signaling. Plant Cell 25:2618–32 [Google Scholar]
  119. Turner AS, Beales J, Faure S, Dunford RP, Laurie DA. 119.  2005. The pseudo-response regulator Ppd-H1 provides adaptation to photoperiod in barley. Science 310:1031–34 [Google Scholar]
  120. Turner AS, Faure S, Zhang Y, Laurie DA. 120.  2013. The effect of day-neutral mutations in barley and wheat on the interaction between photoperiod and vernalization. Theor. Appl. Genet. 126:2267–77 [Google Scholar]
  121. Valverde F, Mouradov A, Soppe W, Ravenscroft D, Samach A, Coupland G. 121.  2004. Photoreceptor regulation of CONSTANS protein in photoperiodic flowering. Science 303:1003–6 [Google Scholar]
  122. Vince-Prue D. 122.  1975. Photoperiodism in Plants London: McGraw-Hill
  123. Wang ZY, Tobin EM. 123.  1998. Constitutive expression of the CIRCADIAN CLOCK ASSOCIATED 1 (CCA1) gene disrupts circadian rhythms and suppresses its own expression. Cell 93:1207–17 [Google Scholar]
  124. Wei X, Xu J, Guo H, Jiang L, Chen S. 124.  et al. 2010. DTH8 suppresses flowering in rice, influencing plant height and yield potential simultaneously. Plant Physiol. 153:1747–58 [Google Scholar]
  125. Wenkel S, Turck F, Singer K, Gissot L, Le Gourrierec J. 125.  et al. 2006. CONSTANS and the CCAAT box binding complex share a functionally important domain and interact to regulate flowering of Arabidopsis. Plant Cell 18:2971–84 [Google Scholar]
  126. Wilhelm EP, Turner AS, Laurie DA. 126.  2009. Photoperiod insensitive Ppd-A1a mutations in tetraploid wheat (Triticum durum Desf.). Theor. Appl. Genet. 118:285–94 [Google Scholar]
  127. Woods DP, Ream TS, Minevich G, Hobert O, Amasino RM. 127.  2014. PHYTOCHROME C is an essential light receptor for photoperiodic flowering in the temperate grass, Brachypodium distachyon. Genetics 198:397–408 [Google Scholar]
  128. Xu Q, Saito H, Hirose I, Katsura K, Yoshitake Y. 128.  et al. 2014. The effects of the photoperiod-insensitive alleles, se13, hd1 and ghd7, on yield components in rice. Mol. Breed. 33:813–19 [Google Scholar]
  129. Xue W, Xing Y, Weng X, Zhao Y, Tang W. 129.  et al. 2008. Natural variation in Ghd7 is an important regulator of heading date and yield potential in rice. Nat. Genet. 40:761–67 [Google Scholar]
  130. Yan L, Loukoianov A, Blechl A, Tranquilli G, Ramakrishna W. 130.  et al. 2004. The wheat VRN2 gene is a flowering repressor down-regulated by vernalization. Science 303:1640–64 [Google Scholar]
  131. Yan L, Loukoianov A, Tranquilli G, Helguera M, Fahima T, Dubcovsky J. 131.  2003. Positional cloning of the wheat vernalization gene VRN1. PNAS 100:6263–68 [Google Scholar]
  132. Yan WH, Wang P, Chen HX, Zhou HJ, Li QP. 132.  et al. 2011. A major QTL, Ghd8, plays pleiotropic roles in regulating grain productivity, plant height, and heading date in rice. Mol. Plant 4:319–30 [Google Scholar]
  133. Yang J, Lee S, Hang R, Kim SR, Lee YS. 133.  et al. 2013. OsVIL2 functions with PRC2 to induce flowering by repressing OsLFL1 in rice. Plant J. 73:566–78 [Google Scholar]
  134. Yokoo T, Saito H, Yoshitake Y, Xu Q, Asami T. 134.  et al. 2014. Se14, encoding a JmjC domain-containing protein, plays key roles in long-day suppression of rice flowering through the demethylation of H3K4me3 of RFT1. PLOS ONE 9:e96064 [Google Scholar]
  135. Zhao J, Huang X, Ouyang X, Chen W, Du A. 135.  et al. 2012. OsELF3-1, an ortholog of Arabidopsis EARLY FLOWERING 3, regulates rice circadian rhythm and photoperiodic flowering. PLOS ONE 7:e43705 [Google Scholar]
  136. Zheng B, Biddulph B, Li D, Kuchel H, Chapman S. 136.  2013. Quantification of the effects of VRN1 and Ppd-D1 to predict spring wheat (Triticum aestivum) heading time across diverse environments. J. Exp. Bot. 64:3747–61 [Google Scholar]
  137. Zuo Z, Liu H, Liu B, Liu X, Lin C. 137.  2011. Blue light-dependent interaction of CRY2 with SPA1 regulates COP1 activity and floral initiation in Arabidopsis. Curr. Biol. 21:841–47 [Google Scholar]
/content/journals/10.1146/annurev-arplant-043014-115555
Loading
/content/journals/10.1146/annurev-arplant-043014-115555
Loading

Data & Media loading...

  • Article Type: Review Article
This is a required field
Please enter a valid email address
Approval was a Success
Invalid data
An Error Occurred
Approval was partially successful, following selected items could not be processed due to error