W box gene transcriptions

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Editor-In-Chief: Henry A. Hoff

File:Arabis neglecta a1.jpg
Arabis neglecta habitat is Tatra Mountains, Orla Perć, Kozia Przełęcz. Credit: Jerzy Opioła.{{free media}}

"Consistent with its role as a transcriptional repressor, OsWRKY71 binds specifically to functionally defined TGAC-containing W boxes of the Amy32b promoter in electrophoretic mobility shift assays (Zhang et al. 2004)."[1]

"Details of auto-regulation or cross-regulation by WRKY factors were provided for the parsley group I member PcWRKY1 and its ortholog AtWRKY33 [37**,39,41]. In response to PAMP treatment PcWRKY1 transcripts accumulate rapidly and transiently [42]. AtWRKY33 is activated with similar kinetics by defense-related stimuli [18,34,41]. This rapid response is mediated by a conserved arrangement of three synergistically acting W boxes (WABC). Chromatin immunoprecipitation (ChIP) revealed that in vivo these orthologous W boxes are constitutively occupied by WRKY proteins [37**,41]."[2]

"The synthesis of SA and the expression of NPR1, a key regulator of some PAMP-triggered responses, appear to be partly controlled by WRKY factors. NPR1 is regulated by WRKY TFs interacting with two W box elements in its 50UTR [44]. Defense-associated SA production is strongly dependent on pathogen-inducible expression of ICS1 [45]. This gene is a likely target of WRKY TFs, as its promoter is enriched for W boxes."[2]

A "missense mutation within its WRKY domain results in conditional activation of defense responses and loss of in vitro binding to W boxes suggesting a negative role of this factor in defense signaling [49]."[2]

Functionality and conservation of the W-box element across plant species shown by gel shift experiments, random binding site selection, yeast one-hybrid screens and co-transfection assays performed with many different WRKY proteins and In silico-based studies have identified clusters of W-boxes in stress-inducible promoters, where the binding of WRKY proteins to W-boxes is a feature of both biotic and abiotic stress responses, together with other plant processes such as germination.[3] It has also been shown that multiple W-boxes have a synergistic effect on transcription.[3]

Almost all WRKY transcription factors bind preferentially to W-boxes, and since their discovery, this has raised the question as to how they show specificity for the promoters of their target genes.[4] Although the W-box core is required, adjacent sequences also play a role in determining binding-site preference.[5] Recent evidence suggests that the TGAC core is more degenerate, composed of a guanine adenine cytosine (GAC) core, and the upstream thymine and downstream pyrimidine flanking sequences help dictate recognition by specific WRKY factors.[6] Basic residues of the WRKY protein domain also are believed to recognize the phosphate backbone of the cis-element.[6]

The solution structure of the C-terminal WRKY domain of Arabidopsis WRKY4 in complex with the W-box DNA has been determined by NMR.[7] A four-stranded β-sheet enters the major groove of DNA in a structure called the β-wedge, where the sheet is nearly perpendicular to the DNA helical ais: as predicted amino acids in the conserved WRKYGQK signature motif contact the W-box DNA.[7]

WRKY

WRKY family of genes contain the common degenerate nucleotide sequence: WRKY, or W = A or T (U) [Weak], R = A or G [puRine], K = G or T (U) [Keto], Y = C or T (U) [pYrimidine].

"Plant immune responses are associated with the concerted modulation of a large number of different WRKY transcripts and proteins [15,34–36,37**]. Upon triggering of SA-dependent defenses, at least 49 AtWRKY genes exhibited differential regulation representing separate waves of transcript accumulation or repression [34]. Their promoters are statistically enriched for W boxes, suggesting that they are autoregulated or controlled by other WRKY proteins [34]."[2]

"Some architectural features of the WRKY web are emerging. As motif D containing group I WRKY TFs can be phosphorylated by MAP-kinases, they are likely to serve as the first WRKY proteins activated in response to PAMP-triggered MAPK signaling. Their targets may include the IIe WRKY genes AtWRKY22 and AtWRKY29, which are upregulated by a PAMP-induced MAPK cascade and contain multiple W boxes within their respective promoters [4]."[2]

Consensus sequences

The "presence of WRKY TF binding sites (C/TTGACC/T, W boxes) in numerous co-regulated Arabidopsis defense gene promoters provided circumstantial evidence that zinc-finger-type WRKY factors play a broad and pivotal role in regulating defenses [10]."[2]

The W box is a DNA cis-regulatory element sequence, (T)TGAC(C/T), which is recognized by the family of WRKY transcription factors.[8][4]

W box samplings

For the Basic programs (starting with SuccessablesWbox.bas) written to compare nucleotide sequences with the sequences on either the template strand (-), or coding strand (+), of the DNA, in the negative direction (-), or the positive direction (+), including extending the number of nts from 958 to 4445, the programs are, are looking for, and found:

  1. negative strand in the negative direction (from ZSCAN22 to A1BG) is SuccessablesWbox--.bas, looking for 3'-(C/T)TGAC(C/T)-5', 5, 3'-CTGACT-5', 17, 3'-TTGACT-5', 130, 3'-TTGACT-5', 307, 3'-CTGACT-5', 1935, 3'-CTGACC-5', 3749,
  2. positive strand in the negative direction is SuccessablesWbox+-.bas, looking for 3'-(C/T)TGAC(C/T)-5', 1, 3'-CTGACC-5', 734,
  3. negative strand in the positive direction (from ZNF497 to A1BG) is SuccessablesWbox-+.bas, looking for 3'-(C/T)TGAC(C/T)-5', 6, 3'-CTGACC-5', 1662, 3'-CTGACC-5', 2213, 3'-TTGACC-5', 2873, 3'-CTGACT-5', 2945, 3'-TTGACC-5', 4018, 3'-CTGACC-5', 4216,
  4. positive strand in the positive direction is SuccessablesWbox++.bas, looking for 3'-(C/T)TGAC(C/T)-5', 3, 3'-TTGACC-5', 1953, 3'-CTGACT-5', 2674, 3'-TTGACT-5', 3735,
  5. complement, negative strand, negative direction is SuccessablesWboxc--.bas, looking for 3'-(A/G)ACTG(A/G)-5', 1, 3'-GACTGG-5', 734,
  6. complement, negative strand, positive direction is SuccessablesWboxc-+.bas, looking for 3'-(A/G)ACTG(A/G)-5', 3, 3'-AACTGG-5', 1953, 3'-GACTGA-5', 2674, 3'-AACTGA-5', 3735,
  7. complement, positive strand, negative direction is SuccessablesWboxc+-.bas, looking for 3'-(A/G)ACTG(A/G)-5', 5, 3'-GACTGA-5', 17, 3'-AACTGA-5', 130, 3'-AACTGA-5', 307, 3'-GACTGA-5', 1935, 3'-GACTGG-5', 3749,
  8. complement, positive strand, positive direction is SuccessablesWboxc++.bas, looking for 3'-(A/G)ACTG(A/G)-5', 6, 3'-GACTGG-5', 1662, 3'-GACTGG-5', 2213, 3'-AACTGG-5', 2873, 3'-GACTGA-5', 2945, 3'-AACTGG-5', 4018, 3'-GACTGG-5', 4216,
  9. inverse complement, negative strand, negative direction is SuccessablesWboxci--.bas, looking for 3'-(A/G)GTCA(A/G)-5', 2, 3'-GGTCAG-5', 1353, 3'-GGTCAA-5', 4416,
  10. inverse complement, negative strand, positive direction is SuccessablesWboxci-+.bas, looking for 3'-(A/G)GTCA(A/G)-5', 6, 3'-AGTCAG-5', 2101, 3'-GGTCAG-5', 2221, 3'-AGTCAG-5', 2608, 3'-AGTCAA-5', 2614, 3'-AGTCAG-5', 2619, 3'-GGTCAG-5', 4270,
  11. inverse complement, positive strand, negative direction is SuccessablesWboxci+-.bas, looking for 3'-(A/G)GTCA(A/G)-5', 6, 3'-GGTCAG-5', 440, 3'-GGTCAG-5', 577, 3'-GGTCAG-5', 713, 3'-GGTCAG-5', 2249, 3'-GGTCAG-5', 2586, 3'-GGTCAA-5', 4308,
  12. inverse complement, positive strand, positive direction is SuccessablesWboxci++.bas, looking for 3'-(A/G)GTCA(A/G)-5', 6, 3'-GGTCAG-5', 2025, 3'-AGTCAG-5', 2099, 3'-GGTCAG-5', 2606, 3'-GGTCAG-5', 2997, 3'-GGTCAG-5', 3083, 3'-GGTCAA-5', 3380,
  13. inverse, negative strand, negative direction, is SuccessablesWboxi--.bas, looking for 3'-(C/T)CAGT(C/T)-5', 6, 3'-CCAGTC-5', 440, 3'-CCAGTC-5', 577, 3'-CCAGTC-5', 713, 3'-CCAGTC-5', 2249, 3'-CCAGTC-5', 2586, 3'-CCAGTT-5', 4308,
  14. inverse, negative strand, positive direction, is SuccessablesWboxi-+.bas, looking for 3'-(C/T)CAGT(C/T)-5', 6, 3'-CCAGTC-5', 2025, 3'-TCAGTC-5', 2099, 3'-CCAGTC-5', 2606, 3'-CCAGTC-5', 2997, 3'-CCAGTC-5', 3083, 3'-CCAGTT-5', 3380,
  15. inverse, positive strand, negative direction, is SuccessablesWboxi+-.bas, looking for 3'-(C/T)CAGT(C/T)-5', 2, 3'-CCAGTC-5', 1353, 3'-CCAGTT-5', 4416,
  16. inverse, positive strand, positive direction, is SuccessablesWboxi++.bas, looking for 3'-(C/T)CAGT(C/T)-5', 6, 3'-TCAGTC-5', 2101, 3'-CCAGTC-5', 2221, 3'-TCAGTC-5', 2608, 3'-TCAGTT-5', 2614, 3'-TCAGTC-5', 2619, 3'-CCAGTC-5', 4270.

Wbox (4560-2846) UTRs

  1. Negative strand, negative direction: CTGACC at 3749.
  2. Negative strand, negative direction: GGTCAA at 4416.
  3. Positive strand, negative direction: GGTCAA at 4308.

Wbox positive direction (4445-4265) core promoters

  1. Negative strand, positive direction: GGTCAG at 4270.

Wbox positive direction (4265-4050) proximal promoters

  1. Negative strand, positive direction: CTGACC at 4216.

Wbox negative direction (2596-1) distal promoters

  1. Negative strand, negative direction: CTGACT at 1935, TTGACT at 307, TTGACT at 130, CTGACT at 17.
  2. Negative strand, negative direction: GGTCAG at 1353.
  3. Positive strand, negative direction: CTGACC at 734.
  4. Positive strand, negative direction: GGTCAG at 2586, GGTCAG at 2249, GGTCAG at 713, GGTCAG at 577, GGTCAG at 440.

Wbox positive direction (4050-1) distal promoters

  1. Negative strand, positive direction: TTGACC at 4018, CTGACT at 2945, TTGACC at 2873, CTGACC at 2213, CTGACC at 1662.
  2. Negative strand, positive direction: AGTCAG at 2619, AGTCAA at 2614, AGTCAG at 2608, GGTCAG at 2221, AGTCAG at 2101.
  3. Positive strand, positive direction: TTGACT at 3735, CTGACT at 2674, TTGACC at 1953.
  4. Positive strand, positive direction: GGTCAA at 3380, GGTCAG at 3083, GGTCAG at 2997, GGTCAG at 2606, AGTCAG at 2099, GGTCAG at 2025.

W box random dataset samplings

  1. Wboxr0: 6, TTGACC at 4380, TTGACT at 4031, CTGACT at 3270, CTGACC at 1834, CTGACC at 1659, TTGACT at 231.
  2. Wboxr1: 5, TTGACT at 4021, TTGACC at 3051, TTGACT at 2937, CTGACC at 1160, CTGACC at 154.
  3. Wboxr2: 4, CTGACC at 3755, CTGACC at 3580, TTGACT at 1736, TTGACC at 853.
  4. Wboxr3: 4, TTGACT at 4479, CTGACC at 4438, TTGACC at 3102, CTGACT at 1141.
  5. Wboxr4: 7, TTGACT at 2961, TTGACC at 2892, TTGACT at 2539, TTGACC at 2131, CTGACT at 1980, CTGACT at 1922, TTGACC at 344.
  6. Wboxr5: 1, TTGACT at 1914.
  7. Wboxr6: 3, CTGACT at 4233, TTGACC at 2398, TTGACT at 2043.
  8. Wboxr7: 2, CTGACC at 3140, TTGACC at 898.
  9. Wboxr8: 2, TTGACC at 1241, TTGACC at 776.
  10. Wboxr9: 3, CTGACC at 4209, TTGACT at 4205, CTGACT at 3521.
  11. Wboxr0ci: 3, AGTCAA at 4044, GGTCAA at 1985, GGTCAG at 846.
  12. Wboxr1ci: 4, GGTCAA at 4484, AGTCAA at 3855, GGTCAA at 2778, GGTCAG at 1751.
  13. Wboxr2ci: 9, AGTCAA at 4375, GGTCAG at 3906, AGTCAA at 3850, GGTCAA at 3164, AGTCAG at 2900, GGTCAA at 2186, GGTCAG at 468, GGTCAA at 142, GGTCAG at 24.
  14. Wboxr3ci: 4, GGTCAG at 3506, GGTCAG at 2239, GGTCAA at 1052, GGTCAA at 601.
  15. Wboxr4ci: 3, AGTCAA at 3240, AGTCAA at 3119, AGTCAA at 986.
  16. Wboxr5ci: 3, AGTCAA at 3608, AGTCAG at 1920, AGTCAG at 1355.
  17. Wboxr6ci: 6, AGTCAA at 4504, AGTCAA at 4450, GGTCAA at 2737, AGTCAG at 2125, GGTCAA at 1833, AGTCAG at 394.
  18. Wboxr7ci: 1, GGTCAA at 4458.
  19. Wboxr8ci: 4, AGTCAA at 4172, GGTCAA at 3494, GGTCAA at 3434, GGTCAA at 2695.
  20. Wboxr9ci: 7, GGTCAA at 4380, AGTCAA at 3950, AGTCAG at 3359, GGTCAA at 3056, AGTCAA at 2475, GGTCAG at 2057, AGTCAG at 107.

Wboxr arbitrary (evens) (4560-2846) UTRs

  1. Wboxr0: TTGACC at 4380, TTGACT at 4031, CTGACT at 3270.
  2. Wboxr2: CTGACC at 3755, CTGACC at 3580.
  3. Wboxr6: CTGACT at 4233.
  4. Wboxr4: TTGACT at 2961, TTGACC at 2892.
  5. Wboxr0ci: AGTCAA at 4044.
  6. Wboxr2ci: AGTCAA at 4375, GGTCAG at 3906, AGTCAA at 3850, GGTCAA at 3164, AGTCAG at 2900.
  7. Wboxr4ci: AGTCAA at 3240, AGTCAA at 3119.
  8. Wboxr6ci: AGTCAA at 4504, AGTCAA at 4450.
  9. Wboxr8ci: AGTCAA at 4172, GGTCAA at 3494, GGTCAA at 3434.

Wboxr alternate (odds) (4560-2846) UTRs

  1. Wboxr1: TTGACT at 4021, TTGACC at 3051, TTGACT at 2937.
  2. Wboxr3: TTGACT at 4479, CTGACC at 4438, TTGACC at 3102.
  3. Wboxr7: CTGACC at 3140.
  4. Wboxr9: CTGACC at 4209, TTGACT at 4205, CTGACT at 3521.
  5. Wboxr1ci: GGTCAA at 4484, AGTCAA at 3855.
  6. Wboxr3ci: GGTCAG at 3506.
  7. Wboxr5ci: AGTCAA at 3608.
  8. Wboxr7ci: GGTCAA at 4458.
  9. Wboxr9ci: GGTCAA at 4380, AGTCAA at 3950, AGTCAG at 3359, GGTCAA at 3056.

Wboxr arbitrary positive direction (odds) (4445-4265) core promoters

  1. Wboxr3: CTGACC at 4438.
  2. Wboxr9ci: GGTCAA at 4380.

Wboxr alternate positive direction (evens) (4445-4265) core promoters

  1. Wboxr0: TTGACC at 4380.
  2. Wboxr2ci: AGTCAA at 4375.

Wboxr arbitrary negative direction (evens) (2811-2596) proximal promoters

  1. Wboxr6ci: GGTCAA at 2737.
  2. Wboxr8ci: GGTCAA at 2695.

Wboxr alternate negative direction (odds) (2811-2596) proximal promoters

  1. Wboxr1ci: GGTCAA at 2778.

Wboxr arbitrary positive direction (odds) (4265-4050) proximal promoters

  1. Wboxr9: CTGACC at 4209, TTGACT at 4205.

Wboxr alternate positive direction (evens) (4265-4050) proximal promoters

  1. Wboxr6: CTGACT at 4233.
  2. Wboxr8ci: AGTCAA at 4172.

Wboxr arbitrary negative direction (evens) (2596-1) distal promoters

  1. Wboxr0: CTGACC at 1834, CTGACC at 1659, TTGACT at 231.
  2. Wboxr2: TTGACT at 1736, TTGACC at 853.
  3. Wboxr4: TTGACT at 2539, TTGACC at 2131, CTGACT at 1980, CTGACT at 1922, TTGACC at 344.
  4. Wboxr6: TTGACC at 2398, TTGACT at 2043.
  5. Wboxr8: TTGACC at 1241, TTGACC at 776.
  6. Wboxr0ci: GGTCAA at 1985, GGTCAG at 846.
  7. Wboxr2ci: GGTCAA at 2186, GGTCAG at 468, GGTCAA at 142, GGTCAG at 24.
  8. Wboxr4ci: AGTCAA at 986.
  9. Wboxr6ci: AGTCAG at 2125, GGTCAA at 1833, AGTCAG at 394.

Wboxr alternate negative direction (odds) (2596-1) distal promoters

  1. Wboxr1: CTGACC at 1160, CTGACC at 154.
  2. Wboxr3: CTGACT at 1141.
  3. Wboxr5: TTGACT at 1914.
  4. Wboxr7: TTGACC at 898.
  5. Wboxr1ci: GGTCAG at 1751.
  6. Wboxr3ci: GGTCAG at 2239, GGTCAA at 1052, GGTCAA at 601.
  7. Wboxr5ci: AGTCAG at 1920, AGTCAG at 1355.
  8. Wboxr9ci: AGTCAA at 2475, GGTCAG at 2057, AGTCAG at 107.

Wboxr arbitrary positive direction (odds) (4050-1) distal promoters

  1. Wboxr1: TTGACT at 4021, TTGACC at 3051, TTGACT at 2937, CTGACC at 1160, CTGACC at 154.
  2. Wboxr3: TTGACC at 3102, CTGACT at 1141.
  3. Wboxr5: TTGACT at 1914.
  4. Wboxr7: CTGACC at 3140, TTGACC at 898.
  5. Wboxr9: CTGACT at 3521.
  6. Wboxr1ci: AGTCAA at 3855, GGTCAA at 2778, GGTCAG at 1751.
  7. Wboxr3ci: GGTCAG at 3506, GGTCAG at 2239, GGTCAA at 1052, GGTCAA at 601.
  8. Wboxr5ci: AGTCAA at 3608, AGTCAG at 1920, AGTCAG at 1355.
  9. Wboxr9ci: AGTCAA at 3950, AGTCAG at 3359, GGTCAA at 3056, AGTCAA at 2475, GGTCAG at 2057, AGTCAG at 107.

Wboxr alternate positive direction (evens) (4050-1) distal promoters

  1. Wboxr0: TTGACT at 4031, CTGACT at 3270, CTGACC at 1834, CTGACC at 1659, TTGACT at 231.
  2. Wboxr2: CTGACC at 3755, CTGACC at 3580, TTGACT at 1736, TTGACC at 853.
  3. Wboxr4: TTGACT at 2961, TTGACC at 2892, TTGACT at 2539, TTGACC at 2131, CTGACT at 1980, CTGACT at 1922, TTGACC at 344.
  4. Wboxr6: TTGACC at 2398, TTGACT at 2043.
  5. Wboxr8: TTGACC at 1241, TTGACC at 776.
  6. Wboxr0ci: AGTCAA at 4044, GGTCAA at 1985, GGTCAG at 846.
  7. Wboxr2ci: GGTCAG at 3906, AGTCAA at 3850, GGTCAA at 3164, AGTCAG at 2900, GGTCAA at 2186, GGTCAG at 468, GGTCAA at 142, GGTCAG at 24.
  8. Wboxr4ci: AGTCAA at 3240, AGTCAA at 3119, AGTCAA at 986.
  9. Wboxr6ci: GGTCAA at 2737, AGTCAG at 2125, GGTCAA at 1833, AGTCAG at 394.
  10. Wboxr8ci: GGTCAA at 3494, GGTCAA at 3434, GGTCAA at 2695.

Wbox analysis and results

Main article: Complex locus A1BG and ZNF497 § Wboxes

The "presence of WRKY TF binding sites (C/TTGACC/T, W boxes) in numerous co-regulated Arabidopsis defense gene promoters provided circumstantial evidence that zinc-finger-type WRKY factors play a broad and pivotal role in regulating defenses [10]."[2]

Reals or randoms Promoters direction Numbers Strands Occurrences Averages (± 0.1)
Reals UTR negative 3 2 1.5 1.5 ± 0.5 (--2,+-1)
Randoms UTR arbitrary negative 21 10 2.1 2.0 ± 0.1
Randoms UTR alternate negative 19 10 1.9 2.0 ± 0.1
Reals Core negative 0 2 0 0
Randoms Core arbitrary negative 0 10 0 0
Randoms Core alternate negative 0 10 0 0
Reals Core positive 1 2 0.5 0.5 ± 0.5 (-+1,++0)
Randoms Core arbitrary positive 2 10 0.2 0.2
Randoms Core alternate positive 2 10 0.2 0.2
Reals Proximal negative 0 2 0 0
Randoms Proximal arbitrary negative 2 10 0.2 0.15
Randoms Proximal alternate negative 1 10 0.1 0.15
Reals Proximal positive 1 2 0.5 0.5 ± 0.5 (-+1,++0)
Randoms Proximal arbitrary positive 1 10 0.1 0.15
Randoms Proximal alternate positive 2 10 0.2 0.15
Reals Distal negative 11 2 5.5 5.5 ± 0.5 (--5,+-6)
Randoms Distal arbitrary negative 24 10 2.4 1.9
Randoms Distal alternate negative 14 10 1.4 1.9
Reals Distal positive 19 2 9.5 9.5 ± 0.5 (-+10,++9)
Randoms Distal arbitrary positive 27 10 2.7 3.4 ± 0.7
Randoms Distal alternate positive 41 10 4.1 3.4 ± 0.7

Comparison:

The occurrences of real W-box UTRs are ≤ randoms, cores, proximals and distals are greater than the randoms. This suggests that the real W boxes are likely active or activable.

Acknowledgements

The content on this page was first contributed by: Henry A. Hoff.

Initial content for this page in some instances came from Wikiversity.

See also

References

  1. ↑ Liu-Min Fan, Xiaoyan Feng, Yu Wang and Xing Wang Deng (2007). "Gibberellin Signal Transduction in Rice". Journal of Integrative Plant Biology. 49 (6): 731−741. doi:10.1111/j.1744-7909.2007.00511.x. Retrieved 16 October 2018.
  2. ↑ 2.0 2.1 2.2 2.3 2.4 2.5 2.6 Thomas Eulgem and Imre E Somssich (2007). "Networks of WRKY transcription factors in defense signaling" (PDF). Current Opinion in Plant Biology. 10: 366–371. doi:10.1016/j.pbi.2007.04.020. Retrieved 17 October 2018.
  3. ↑ 3.0 3.1 Rushton, Paul. The Lab of Dr. Paul Rushton. wordpress.com. Retrieved 17 June 2013.
  4. ↑ 4.0 4.1 Rushton PJ, Somssich IE, Ringler P, Shen QJ (May 2010). "WRKY transcription factors". Trends Plant Science. 15 (5): 247–58. doi:10.1016/j.tplants.2010.02.006. PMID 20304701.
  5. ↑ Ciolkowski, I.; Wanke D; Birkenbihl RP; Somssich IE. (2008). "Studies on DNA-binding selectivity of WRKY transcription factors lend structural clues into WRKY-domain function". Plant Mol Biol. 68: 81–92. doi:10.1007/s11103-008-9353-1. PMC 2493524. PMID 18523729.
  6. ↑ 6.0 6.1 Brand; Fischer; Harter; Kohlbacher; Wanke (2013). "Elucidating the evolutionary conserved DNA-binding specificities of WRKY transcription factors by molecular dynamics and in vitro binding assays". Nucleic Acids Research. 41 (21): 9764–9778. doi:10.1093/nar/gkt732.
  7. ↑ 7.0 7.1 Yamasaki, K.; Kigawa T; Watanabe S; Inoue M; Yamasaki T; Seki M; Shinozaki K; Yokoyama S. (2012). "Structural basis for sequence-specific DNA recognition by an Arabidopsis WRKY transcription factor". J. Biol. Chem. 287: 7683–91. doi:10.1074/jbc.M111.279844. PMC 3293589. PMID 22219184.
  8. ↑ Rushton, Paul; Macdonald, H.; Huttly, A.K.; Lazarus, C.M.; Hooley, R (1995). "Members of a new family of DNA-binding proteins bind to a conserved cis-element in the promoters of alpha-Amy2 genes". Plant Molecular Biology. 29: 691–702. doi:10.1007/bf00041160. PMID 8541496.

External links

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