Figures (2)  Tables (3)
    • Figure 1. 

      Functions of grape WRKY transcription factors in biotic stresses. The expression of grape WRKY members were induced (solid blue arrows) by pathogen attack (DM, Plasmopara viticola; PM, Erysiph ecichoraceaum; WR, Coniella diplodiella), hormone (JA, Methyl jasmonate; SA, Salicylic acid; Et, Ethylene), and wound. Grape WRKY members enhanced (solid arrows) or repressed (dotted lines) plant resistance to different types of pathogens. Grape WRKY members in red font were involved in regulating the expression of JA signaling pathway-related denfense genes. Grape WRKY members in black font were involved in regulating the expression of SA signaling pathway-related defence genes. Grape WRKY members involved in regulating the expression of JA signaling pathway-related defence genes (red font), SA signaling pathway-related defence genes (black font), both SA and JA signaling pathway-related defence genes (blue font), or unknown (purple font). The function of WRKY members were identified in grapevines (red arrows) or other plants (black arrows).

    • Figure 2. 

      Functions of grape WRKY transcription factors in abiotic stresses. The expression of grape WRKY members were induced (solid blue arrows) by abiotic stress, and hormones (ABA, Abscisic acid; Et, Ethylene). Grape WRKY members were involved in plant hormone synthesis (red dotted arrows). Grape WRKY members enhanced (solid arrows) or repressed (dotted arrows) plant abiotic stresses tolerance.

    • Gene nameGene numberLocationGene numberFormer names
      VvWRKY1GSVIVT01012196001chr1:628,681..633,595VIT_01s0011g00720
      VvWRKY2GSVIVT01020060001chr1:10,977,206..10,982,627VIT_01s0026g01730
      VvWRKY3GSVIVT01010525001chr1:21,460,152..21,461,423VIT_01s0010g03930VlWRKY3[41], VvWRKY8[51], VvWRKY3[49]
      VvWRKY4GSVIVT01019419001chr2:512,179..513,535VIT_02s0025g00420
      VvWRKY5GSVIVT01019511001chr2:1,228,115..1,229,951VIT_02s0025g01280
      VvWRKY6GSVIVT01035426001chr4:1,210,142..1,211,252VIT_04s0008g01470
      VvWRKY7GSVIVT01035884001chr4:5,247,432..5,248,815VIT_04s0008g05750
      VvWRKY8GSVIVT01035885001chr4:5,265,887..5,267,902VIT_04s0008g05760VpWRKY3[59]
      VvWRKY9GSVIVT01035965001chr4:6,569,931..6,576,445VIT_04s0008g06600
      VvWRKY10GSVIVT01033188001chr4:9,363,044..9,365,026VIT_04s0069g00920VvWRKY11[17]
      VvWRKY11GSVIVT01033194001chr4:9,399,822..9,400,803VIT_04s0069g00970
      VvWRKY12GSVIVT01033195001chr4:9,409,805..9,411,286VIT_04s0069g00980
      VvWRKY13GSVIVT01019109001chr4:16,664,476..16,666,517VIT_04s0023g00470
      VvWRKY14GSVIVT01034968001chr5:530,097..531,696VIT_05s0077g00730
      VvWRKY15GSVIVT01025491001chr6:364,396..365,481VIT_06s0004g00230
      VvWRKY16GSVIVT01024624001chr6:8,290,089..8,292,826VIT_06s0004g07500VaWRKY33[57]
      VvWRKY17GSVIVT01000752001chr7:381,054..383,675VIT_07s0141g00680
      VvWRKY18GSVIVT01028129001chr7:4,044,185..4,045,807VIT_07s0005g01520
      VvWRKY19GSVIVT01028147001chr7:4,200,461..4,202,183VIT_07s0005g01710
      VvWRKY20GSVIVT01028244001chr7:4,899,940..4,903,113VIT_07s0005g02570
      VvWRKY21GSVIVT01022067001chr7:16,322,554..16,324,062VIT_07s0031g00080
      VvWRKY22GSVIVT01022245001chr7:17,794,621..17,797,164VIT_07s0031g01710
      VvWRKY23GSVIVT01022259001chr7:17,958,339..17,960,800VIT_07s0031g01840VvWRKY13[28,33,52,53]
      VvWRKY24GSVIVT01030258001chr8:9,796,097..9,798,804VIT_08s0058g00690VvWRKY33[29], VvWRKY24[49]
      VvWRKY25GSVIVT01030174001chr8:10,843,800..10,846,082VIT_08s0058g01390VpWRKY1[16]
      VvWRKY26GSVIVT01025562001chr8:14,033,449..14,039,296VIT_08s0040g03070VvWRKY26[34]
      VvWRKY27GSVIVT01034148001chr8:14,828,036..14,830,056VIT_08s0007g00570
      VvWRKY28GSVIVT01015952001chr9:16,094,245..16,096,198VIT_09s0018g00240VaWRKY14[55], VvWRKY40[30]
      VvWRKY29GSVIVT01012682001chr10:618,665..621,093VIT_10s0116g01200VaWRKY12[20], VpWRKY31[45]
      VvWRKY30GSVIVT01021252001chr10:3,008,699..3,010,258VIT_10s0003g01600
      VvWRKY31GSVIVT01021397001chr10:4,894,494..4,896,041VIT_10s0003g02810
      VvWRKY32GSVIVT01021765001chr10:10,756,001..10,759,241VIT_10s0003g05740
      VvWRKY33GSVIVT01023600001chr11:7,838,041..7,845,641VIT_11s0037g00150VpWRKY2[16]
      VvWRKY34GSVIVT01029265001chr11:17,821,900..17,823,218VIT_11s0052g00450
      VvWRKY35GSVIVT01020864001chr12:878,978..881,517VIT_12s0028g00270
      VvWRKY36chr12:2,348,148..2,350,455XP_002277383.1
      VvWRKY37GSVIVT01030453001chr12:5,678,804..5,681,082VIT_12s0059g00880
      VvWRKY38GSVIVT01030046001chr12:9,116,720..9,122,740VIT_12s0057g00550
      VvWRKY39GSVIVT01029688001chr12:13,065,135..13,067,116VIT_12s0055g00340
      VvWRKY40GSVIVT01032662001chr13:1,716,836..1,718,877VIT_13s0067g03130
      VvWRKY41GSVIVT01032661001chr13:1,719,393..1,720,803VIT_13s0067g03140
      VvWRKY42GSVIVT01036223001chr14:8,753,748..8,755,749VIT_14s0081g00560
      VvWRKY43GSVIVT01033063001chr14:25,479,103..25,481,917VIT_14s0068g01770
      VvWRKY43[49]
      VvWRKY44GSVIVT01011356001chr14:28,924,779..28,926,499VIT_14s0108g00120
      VvWRKY45GSVIVT01011472001chr14:29,916,401..29,920,389VIT_14s0108g01280
      VvWRKY46GSVIVT01018300001chr15:11,499,221..11,503,179VIT_15s0021g01310
      VvWRKY47GSVIVT01027069001
      chr15:18,191,021..18,193,470
      VIT_15s0046g01140MrWRKY30[31], VlWRKY48[42], VvWRKY46[19,44]
      VvWRKY48GSVIVT01026969001chr15:18,940,954..18,942,146VIT_15s0046g02150
      VvWRKY49GSVIVT01026965001chr15:18,957,235..18,958,812VIT_15s0046g02190
      VvWRKY22[26]
      VvWRKY50GSVIVT01028823001chr16:18,360,079..18,360,711VIT_16s0050g01480
      VvWRKY51GSVIVT01028718001


      chr16:19,477,129..19,479,547
      VIT_16s0050g02510VqWRKY52[43], VvWRKY52[44], VvWRKY30[21], VdWRKY53[56]
      VvWRKY52GSVIVT01008553001chr17:922,644..925,087VIT_17s0000g01280VvWRKY1[18,40]
      VvWRKY53GSVIVT01008046001chr17:6,316,168..6,320,034VIT_17s0000g05810VvWRKY53[49]
      VvWRKY54GSVIVT01009441001chr18:8,392,053..8,393,721VIT_18s0001g10030
      VvWRKY55GSVIVT01037686001chr19:6,882,399..6,884,987VIT_19s0090g00840
      VvWRKY56GSVIVT01037775001chr19:7,760,183..7,767,013VIT_19s0090g01720
      VvWRKY57GSVIVT01014854001chr19:10,665,189..10,669,055VIT_19s0015g01870
      VvWRKY58GSVIVT01001332001chr1_random:297,588..311,938VIT_01s0011g00220
      VvWRKY2[37,54,60]
      VvWRKY59GSVIVT01007006001chrUn:29,694,084..29,699,952VIT_00s0463g00010
      VvWRKY60GSVIVT01001286001chr2:4,989,461..4,989,778VIT_02s0154g00210
      VvWRKY61GSVIVT00037762001chr16:18360079..18360711VIT_16s0050g01480

      Table 1. 

      Putative grape WRKY genes identified in the grape genome.

    • Former nameGene function
      VlWRKY3[41]Enhanced resistance to Golovinomyces cichoracearum, but increased susceptibility to Botrytis cinerea in Arabidopsis
      VpWRKY3[59]Enhanced tolerance to Ralstonia solanacearum in tobacco
      VvWRKY33[29]Enhanced resistance to Plasmopara viticola in grapevine; participate in resistance to B. cinerea in Arabidopsis
      VpWRKY1[16]Enhanced resistance to Erysiphe necator in Arabidopsis
      VvWRKY40[30]Increased susceptible to Phytophthora capsica in tobacco
      VpWRKY31[45]Enhanced resistance to E. necator in susceptible V. vinifera
      VpWRKY2[16]Enhanced resistance to E. necator in Arabidopsis
      MrWRKY30[31]Enhanced resistance to Hyaloperonospora parasitica in Arabidopsis
      VlWRKY48[42]Enhanced resistance to G. cichoracearum in Arabidopsis
      VlWRKY46[19]Attenuated phylloxera attack and delayed nymph development in susceptible grape cultivars
      VqWRKY52[43,44]Enhanced resistance to G. cichoracearum and Pseudomonas syringae pv. tomato DC3000, but increased susceptibility to B. cinerea in Arabidopsis. Enhanced resistance to B. cinerea in gene-edited grape
      VdWRKY53[56]Enhanced resistance to Coniella diplodiella, Pst DC3000 and G. cichoracearum in Arabidopsis
      VvWRKY1[18,40]Enhanced resistance to Pythium, Peronospora tabacina, and E. cichoracearum in tobacco. Enhanced resistance to P. viticola in grapevine
      VqWRKY53[50]Enhanced resistance to Pst DC3000 in Arabidopsis

      Table 2. 

      Functional analysis of grape WRKY proteins under biotic stresses.

    • Gene nameGene function
      VlWRKY3[41]Enhanced tolerance to the salt and drought stress in Arabidopsis
      VpWRKY3[59]Enhanced tolerance to the salt stress in tobacco
      VvWRKY11[17]Enhanced tolerance to the osmotic stress in Arabidopsis
      VaWRKY33[57]Enhanced tolerance to the cold stress in Arabidopsis and grape calli
      VvWRKY13[52,53]Negatively regulated drought and salt stress tolerance of Arabidopsis
      VpWRKY1[16]Enhanced tolerance to the salt stress in Arabidopsis
      VaWRKY14[55]Enhanced tolerance to the drought stress in Arabidopsis
      VaWRKY12[20]Enhanced tolerance to the cold stress in Arabidopsis and grape calli
      VpWRKY2[16]Enhanced tolerance to the salt and cold stress in Arabidopsis
      VlWRKY48[42]Enhanced tolerance to the drought stress in Arabidopsis
      MrWRKY30[31]Enhanced tolerance to the cold stress in Arabidopsis, and negatively regulated salt stress tolerance of Arabidopsis
      VvWRKY30[21]Enhanced tolerance to the salt stress in Arabidopsis
      VvWRKY2[54]Enhanced tolerance to the salt stress and osmotic stress in tobacco

      Table 3. 

      Functional analysis of grape WRKY proteins under abiotic stresses.