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Table 1 The detailed information of genes association with the wheat heading-flowering regulatory network

From: Gene regulatory network and abundant genetic variation play critical roles in heading stage of polyploidy wheat

Gene Chromosomr. location Protein product Description Functions Mutant phenotype Gene number in IWGSC1.0 Position in IWGSC1.0 References
Vrn1 5A, 5B, 5D Encodes a MADS-box transcription factor, also named as WAP1 or TaVRT1 Induced by prolonged cold, long photoperiod Not only promotes the apex transition to generative development, but also activation of long day response in leaves Recessive alleles at all Vrn-1 homoeoloci confer a winter growth habit (vernalization sensitive), whereas one or more dominant alleles at Vrn-1 homoeoloci result in a spring growth habit (vernalization insensitive). TraesCS5A01G391700, TraesCS5B01G396600, TraesCS5D01G401500. 5A:587423056–587,423,240, 5B:573815719–573,815,903, 5D:467184094–467,184,278. [16, 22, 23]
Vrn2 5A Encodes a protein containing a putative zinc finger and a CCT protein-protein interaction domain Down-regulated when plants are vernalized A dominant repressor of flowering Functional mutations in the ZCCT genes result in a spring growth habit and early flowering No No [17]
Vrn3 7B Encodes a mobile protein, homologous to the Arabidopsis FLOWERING LOCUS T (FT) Induced by vernalization and long days Accelerates reproductive apex development Transgenic wheat plants overexpressing Vrn-3B have an extra-early flowering phenotype without the need of vernalization TraesCS7B01G013100 7B:9700818–9,704,363 [18]
Vrn4 5D Encodes a MADS-box transcription factor highly similar to VRN1 VRN-D4 locus originated by the insertion of a large segment from chromosome arm 5AL into chromosome arm 5DS Vrn-D4 likely operates upstream of the positive regulatory feedback loop connecting Vrn1, Vrn2 and Vrn3. The mutation flowered later than plants carrying the wild type allele No No [30, 31]
VER2 2D Encodes a nucleocytoplasmic carbohydrate-binding protein, a jacalin-like lectin, with high affinity for glcnac and galactose After vernalization, VER2 accumulates predominantly in the nucleus in shoot tips and young leaves Nuclear-localized VER2 interacts with O-glcnac-modified TaGRP2 to relieve the repression on tavrn1 transcript accumulation and to promote flowering in hexaploid winter wheat VER2 downregulated in winter wheat results in flowered later No No [47, 50]
TaGRP2 No. Glycine-rich RNA binding protein TaGRP2 is dynamically O-glcnacylated during vernalization TaGRP2 binds to the pre-mRNA of Vrn1 and inhibits Vrn1 expression TaGRP2-Ri plants accelerated flowering compared with wild type No No [50, 51]
TaVRT2 7A, 7B, 7D Encodes a predicted protein of 226 amino acids, belongs to the StMADS 11 Accumulate in winter wheat during the vegetative phase and decline towards the transition to the reproductive phase. The presence of Vrn2 and TaVRT-2 transcripts early during long time exposure could reduce or delay the expression of Vrn1 Mutation of TaVRT-2 results in the advance of flowering No No [48]
TaGI 3A, 3B, 3D Encodes a nucleoplasmically localized protein which contains 1174 amino acid residues The patterns of TaGI rhythmic expression in leaves are regulated by circadian clocks, but can be disturbed by light/dark cycle Functions in mediating photoperiodic flowering, controlling circadian rhythms and phytochrome signaling Mutations in the TaGI gene cause delayed flowering only in long day photoperiod TraesCS3A01G116300; TraesCS3B01G135400; TraesCS3D01G118200. 3A:84189859–84,191,364; 3B:117928502–117,930,007; 3D:71969619–71,970,784. [63, 64]
WSOC1 4DL Is a member of the monocot SOC1-like gene family WSOC1 expression is affected neither by vernalization nor photoperiod, whereas it is induced by gibberellin at the seedling stage WSOC1 functions as a flowering activator like SOC1 in Arabidopsis Downregulated of WSOC1 results in delayed flowering TraesCS4D01G341700 4D:498394464–498,398,154 [71]
WPCL1 No. Encodes a MYB transcription factor belonging to the GARP family Maybe regulated by circadian clock Controlling the early flowering phenotype in the einkorn wheat mutant Deletion of WPCL1 leads to flowering even under short-day conditions No No [60, 61]
TaHD1 6A, 6B, 6D Encodes a transcription factor with zinc finger motif and nuclear localization signals, also called CO2 Can regulated by long-day condition and circadian clock Directly regulate vernalization gene under long-day condition The co mutant show a delayed flowering response under long-day environment TraesCS6A01G289400; TraesCS6B01G319500; TraesCS6D01G269500. 6A:521453035–521,453,945; 6B:567398838–567,399,457; 6D:379572052–379,572,962. [44, 62]
TaPHYC 5A, 5B, 5D No Long-day induced wheat PHYC forms signaling active homodimers and translocate into the nucleus Promotes wheat flowering under inductive photoperiods The loss of function of wheat PHYC results in altered expression of circadian clock and photoperiod genes and a dramatic delay in flowering under long days TraesCS5A01G391300; TraesCS5D01G401000; TraesCS5B01G396200. 5A:586595153–586,599,481; 5D:466221190–466,223,373; 5B:573216947–573,219,055 [60, 61]
Ppd1 2A, 2B, 2D Encodes a pseudo-response regulator (PRR) protein with a CCT domain, also named taprr37 Induced by long-day Can regulated vernalization genes and participate in circadian clock function Knockdown of Ppd1 made the wheat delayed flowering TraesCS2A01G081900; TraesCS2D01G079600 2A:36936362–36,938,400; 2D:33952488–33,955,629. [8, 57]