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Research ArticleBIOCHEMICAL PROCESSES AND MACROMOLECULAR STRUCTURES
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Genetic Control of Natural Variation in Arabidopsis Glucosinolate Accumulation

Daniel J. Kliebenstein, Juergen Kroymann, Paul Brown, Antje Figuth, Deana Pedersen, Jonathan Gershenzon, Thomas Mitchell-Olds
Daniel J. Kliebenstein
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Juergen Kroymann
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Paul Brown
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Antje Figuth
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Deana Pedersen
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Jonathan Gershenzon
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Thomas Mitchell-Olds
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Published June 2001. DOI: https://doi.org/10.1104/pp.126.2.811

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  • Fig. 1.
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    Fig. 1.

    Glucosinolate biosynthetic pathway. I, 2-Alkylmalic acid; II, 3-alkylmalic acid; III, 2-oxo acid. A, Basic glucosinolate structure. B, Outline of the pathway which can be divided into three parts: elongation of the amino acid side chain, formation of the basic glucosinolate skeleton, and further side chain modification. Each chain elongation cycle adds an additional methylene group (Graser et al., 2000).

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    Fig. 2.

    Side chain modifications of Met-derived glucosinolates in Arabidopsis. Potential side chain modifications for the elongated Met derivative, C4 dihomo-Met, are shown. Steps with natural variation identified in this study are shown in bold to the right or left of each enzymatic arrow.

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    Fig. 3.

    Effect of GS-OX on conversion of methylthioalkyl to methylsulfinylalkyl glucosinolates. Bar diagrams show the ratio of methylthioalkyl (MT) glucosinolate to methylsulfinylalkyl (MSO) glucosinolate content in leaf (A) and seeds (B) for each ecotype analyzed.

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    Fig. 4.

    GS-AOP regulates the accumulation of aliphatic glucosinolates in the leaves of Arabidopsis. The bars depict the average total aliphatic glucosinolate accumulation in leaves of 39 ecotypes. The ecotypes are classified based on the inferred genotype at three biosynthetic loci: GS-Elong, either C3- or C4-accumulating ecotypes; GS-AOP, alkenyl-, hydroxypropyl-, or methylsulfinylalkyl-containing ecotypes; and GS-OH, functional or nonfunctional alleles.

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    Fig. 5.

    Correlation of glucosinolate accumulation between the leaves and seeds. A, Scatter plot depicting the relationship between aliphatic glucosinolate accumulation in leaves (ALIPHL) and seeds (ALIPHS) of the ecotypes tested. The 90% confidence ellipse interval is drawn for reference. The values are in μmol g dry weight−1. B, Scatter plot depicting the relationship between indolic glucosinolate accumulation in the leaves (INDOLEL) and seeds (INDOLES) of the ecotypes tested. The values are in μmol g dry weight−1.

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    Fig. 6.

    Map of GS-OX on Chromosome I. Ninety-two F2 progeny were scored for the microsatellites indicated and for the GS-OX biochemical phenotype. The distance from the AthGeneA and nga692 markers to GS-OX is shown in cM to the right of the arrows. The approximate location of GS-OX is shown to the left of the chromosome.

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    Fig. 7.

    Correlation of C7 to C8 and C3 to C4 intermediates in the biosynthesis of chain-elongated Met-derived glucosinolates. A, Scatter plot showing the correlation of the conversion of C3 to C4 in the seeds and leaves. B, Scatter plot showing the correlation of the conversion of C7to C8 in the seeds and leaves. C, Scatter plot showing the correlation of the conversion of C3to C4 with C7 to C8 in the leaves. D, Scatter plot showing the correlation of the conversion of C3 to C4 with C7 to C8 in the seeds.

Tables

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    Table I.

    Glucosinolates in the leaves of arabidopsis ecotypes

    EcotypeGenotype1234567811
    EAOHOX
    Ag-04332–––1.55–4.691.568.10
    Bla-104331–––0.400.831.540.190.362.19
    Bs14332–––1.43–4.291.245.65
    Cal4332–––0.32–1.014.9011.99
    Cnt4332––0.312.47–7.170.2812.46
    Ema-14332––0.131.93–5.830.218.28
    Pog-043320.06–0.322.73–7.980.219.39
    Tac4332–0.050.152.27–6.770.357.38
    Kas4322––––––5.3111.64
    Sorbo4322––––––4.5520.96
    Cvi4312––––––10.0324.40
    Di-142–21.790.03––3.62–––
    Aa-041–2––––1.31–0.070.45
    Col41–2–0.37––4.96–0.03–
    Ma-041–2––––2.39–––
    Mt-041–2––––3.98–––
    Can33–1–0.61––––18.430.40
    Kondara33–20.11–––––25.990.47
    Ei-233–2–––––0.0416.790.19
    Hodja33–2––––––13.100.18
    Ita-033–2––––––––
    Kil-033–2––––––14.570.19
    Mr-033–2––––––––
    Mrk-033–20.15–––––20.270.29
    Rsch-033–2––––––10.020.22
    Su-033–1–0.26––––12.010.07
    Wl-033–20.06–––––14.380.16
    Bl-132–27.32–––––0.200.03
    Di-g32–2––––––––
    Ka-032–214.320.04––––––
    Ler32–29.070.17––0.02–––
    Lip-032–25.140.68––––––
    No-032–27.220.34––0.08–––
    Pet32–28.550.69––0.03–––
    Pi-032–22.07–––0.08–––
    Sei-032–23.760.99––0.07–––
    Tsu-132–25.190.62––0.05––0.02
    Yo-032–25.400.49––0.20–––
    Oy-031–2––––––––
    Average––––4.680.410.231.641.364.377.605.44
    sd ––––4.150.300.100.901.782.897.967.08
    Minimum––––0.060.030.130.320.020.040.030.02
    Maximum––––14.320.990.322.734.967.9825.9924.40
    13141516181920212225263032
    ––0.040.210.46–1.113.190.360.020.570.100.71
    –0.840.020.170.054.951.121.700.42–0.430.100.41
    0.02–0.040.220.36–1.072.530.31–0.210.120.36
    0.090.160.030.360.51–1.772.430.470.090.280.210.52
    0.12–0.030.420.830.053.003.140.320.010.390.090.56
    0.10–0.010.340.60–1.843.240.200.010.330.110.42
    0.11–0.030.370.73–1.872.980.310.030.760.060.36
    0.05–0.010.250.470.041.413.160.090.010.330.060.26
    ––0.020.060.090.020.491.490.350.020.960.040.29
    ––0.020.170.30–1.211.730.310.020.450.070.29
    0.09–0.010.290.29–2.241.080.19–0.390.090.38
    0.01–0.040.24–0.511.242.820.440.030.560.050.20
    ––0.020.11–0.531.081.870.28–0.04–0.21
    ––0.020.16–0.970.992.880.31–0.620.030.19
    ––0.020.10–0.150.861.730.32–0.52–0.07
    ––0.050.15–0.301.132.810.52–0.78–0.15
    –3.660.010.18–0.271.772.140.47–0.380.151.00
    –0.04–0.11––1.892.620.42–1.010.030.53
    ––0.010.12––1.762.330.53–0.480.080.86
    ––––––0.611.160.16–0.490.030.30
    –––––––––––––
    –0.060.010.14––1.822.520.25–0.340.080.58
    –––––––––––––
    ––0.010.12––2.122.090.16–1.070.060.56
    –––0.09––1.341.880.19–0.710.010.24
    –2.020.010.07–0.061.361.970.25–0.640.060.79
    0.020.010.010.150.05–1.222.000.18–0.780.020.18
    –0.080.020.10–0.051.681.760.33–0.940.030.67
    –––––––––––––
    ––0.010.08––1.541.780.26–0.900.040.45
    ––0.020.07––1.321.940.21–0.710.030.33
    –0.040.010.08––1.041.730.33–0.760.020.27
    –0.090.030.03–0.030.881.770.18–0.320.040.52
    –0.140.030.08–0.061.692.820.26–1.130.040.50
    –––0.04––0.640.930.05–0.46––
    –0.020.050.10––1.112.060.06–0.38–0.07
    –0.210.030.09–0.041.132.770.15–0.510.050.40
    –0.120.020.07–0.010.812.910.11–0.770.040.35
    –––––––––––––
    0.070.540.020.160.400.501.382.230.280.030.580.060.41
    0.041.050.010.100.261.220.520.630.130.030.260.040.22
    0.010.010.010.030.050.010.490.930.050.010.040.010.07
    0.123.660.050.420.834.953.003.240.530.091.130.211.00
    Totals
    AliphIndoleMTMSOAOPC3C4C7C8C4PerC8Per
    18.494.160.811.3216.361.5614.340.311.820.9020.854
    13.152.576.302.124.371.039.910.271.530.9060.850
    14.763.090.481.3112.971.2411.370.341.430.9020.808
    21.843.210.892.2218.735.0613.320.572.290.7250.801
    27.763.880.703.5423.520.2822.460.513.560.9880.875
    19.793.780.532.2816.980.2116.170.452.260.9870.834
    24.194.080.422.3521.420.2720.420.432.230.9870.838
    19.513.590.361.7617.390.4016.610.311.670.9670.843
    17.942.820.350.5517.045.3111.660.100.780.6870.886
    27.552.510.361.3825.814.5520.960.241.500.8220.862
    37.811.670.472.6234.7210.0324.400.382.620.7090.873
    7.693.860.765.141.791.824.130.291.440.6940.832
    3.762.210.742.500.520.072.290.111.290.9700.921
    7.703.831.196.480.030.405.930.191.180.9370.861
    3.572.590.223.350.000.002.540.100.931.0000.903
    5.714.160.455.260.000.004.280.151.281.0000.895
    26.473.005.082.5618.8322.700.670.332.770.0290.894
    29.174.050.602.0026.5726.140.470.142.420.0180.945
    19.843.350.941.8817.0216.790.230.202.620.0140.929
    14.221.810.330.6113.2813.100.180.030.910.0140.968
    ––––––––––
    17.443.120.721.9614.7614.630.190.222.400.0130.916
    –––––––––––
    23.573.330.622.2420.7120.420.290.182.680.0140.937
    11.922.780.251.4310.2410.020.220.101.580.0210.940
    16.702.872.931.6912.0814.290.130.132.150.0090.943
    16.252.970.211.3914.6514.450.160.171.400.0110.892
    10.163.050.831.787.557.600.080.132.350.0100.948
    –––––––––––
    16.472.950.491.6614.3214.360.000.121.990.0000.943
    11.012.880.361.589.079.240.020.101.650.0020.943
    7.272.830.331.805.145.860.000.101.310.0000.929
    9.232.300.681.337.227.650.110.071.400.0140.952
    11.784.240.742.498.559.380.090.122.190.0100.948
    2.831.440.000.762.072.070.080.040.640.0370.841
    6.122.550.092.273.764.770.070.101.180.0140.922
    7.803.460.701.895.216.020.110.141.530.0180.916
    7.493.810.521.575.406.010.210.111.160.0340.913
    –––––––––––
    15.343.110.902.2012.237.365.830.211.780.413–
    8.430.731.301.268.667.107.920.140.660.448–
    2.831.440.000.550.000.000.000.030.640.000–
    37.814.246.306.4834.7226.1424.400.573.561.000–

    Quantities are given in μmol g dry wt−1 and are the mean of three extractions of each ecotype. The nos. at the top of each column refer to the list of glucosinolate in “Materials and Methods.” The values underneath the totals section are as follows: Aliph, sum of aliphatic glucosinolates; Indole, sum of indolic glucosinolates; MT, sum of methylthio glucosinolates; MSO, sum of methylsulfinyl glucosinolates; AOP, sum of alkenyl and hydroxy aliphatic glucosinolates; C3, sum of three carbon aliphatic glucosinolates; C4, sum of four carbon aliphatic glucosinolates; C7, sum of seven carbon aliphatic glucosinolates; C8, sum of eight carbon aliphatic glucosinolates; and C4 per, C4/(C3 + C4), C8per = C8/(C7 + C8).

      • View popup
      Table II.

      Glucosinolates in the seeds of Arabidopsis ecotypes

      Genotypes1234567
      EAOHOXOHB
      Ag-043322––1.02––2.400.15
      Bla-1043312––0.41––1.12–
      Bs1433220.31–1.182.02–8.170.29
      Cal43321–––––0.460.21
      Cnt43321–––4.20–13.660.30
      Ema-143322––0.580.65–2.38–
      Pog-043321–––2.91–7.67–
      Tac433210.190.15–2.47–5.840.13
      Kas432220.46–0.490.42–1.430.98
      Sorbo432220.37–0.890.25–1.58–
      Cvi431210.36–––––0.96
      Di-142–210.93–––0.74–0.12
      Aa-041–220.26–0.81–0.52––
      Col41–220.69–2.20–1.87––
      Ma-041–220.47–1.11–0.22–0.21
      Mt-041–220.37–1.43–0.71––
      Can33–1–0.39–––––1.64
      Kondara33–2–4.190.90––––6.96
      Ei-233–2–2.53–––––4.44
      Hodja33–2–2.271.41––––7.60
      Ita-033–2–1.831.40––––1.36
      Kil-033–2–1.360.55––––2.50
      Mr-033–2–0.84––––0.131.63
      Mrk-033–2–1.470.80––––1.54
      Rsch-033–2––––––––
      Su-033–1–0.71––0.13–0.640.99
      Wl-033–2–2.170.33––––3.87
      Bl-132–2–1.88––––––
      Di-g32–2–4.17––––––
      Ka-032–2–10.91–––––0.29
      Ler32–2–7.860.19––––0.12
      Lip-032–2–3.26––––––
      No-032–2–14.160.50––0.28––
      Pet32–2–8.47––––––
      Pi-032–2–3.24––––––
      Sei-032–2–2.580.20–––––
      Tsu-132–2–3.615.58–––––
      Yo-032–2–2.56–––––0.16
      Oy-031–2–1.230.62–––––
      Average–––––2.691.051.011.630.723.791.66
      sd –––––3.291.490.531.500.604.182.18
      Minimum–––––0.190.150.410.130.220.130.12
      Maximum–––––14.165.582.204.201.8713.667.60
      91011121314151617181920
      –0.250.47–0.07––0.56–0.261.724.75
      –0.160.14–0.180.80–0.280.180.073.742.47
      –0.340.52––––0.33–0.116.862.02
      ––0.470.250.05––0.510.110.171.953.41
      –0.492.870.150.15––0.85–0.582.264.55
      –0.460.17–0.25–0.020.92–0.221.526.01
      –0.550.62–0.100.08–0.77–0.292.935.99
      –0.420.650.10–0.23–0.50–0.284.664.56
      ––1.51––0.100.030.22–0.070.612.99
      ––12.52––0.120.020.34–0.313.342.72
      ––3.310.13–0.09–0.32–0.134.014.53
      ––––0.070.30–0.65––13.095.28
      ––––––0.020.22––11.701.90
      0.19–––0.070.080.060.55––18.952.76
      –––––0.220.030.16––7.662.49
      –––––0.100.010.31––12.742.49
      –––––0.15–0.290.06–0.266.81
      ––0.14––3.20–0.15–0.070.273.29
      –––––0.89–0.19––0.174.80
      ––0.13––6.850.020.11––0.262.61
      –––––9.04–0.34–0.150.726.43
      –––––3.33–0.28––0.116.07
      –––––0.660.020.28––0.258.19
      –––––5.90–0.28––0.365.83
      ––––––––––––
      –––0.10–0.940.020.17––0.484.17
      –––––4.07–0.12––0.212.46
      –––––2.090.020.16––0.484.60
      –––––0.69–0.20––0.414.93
      –––––0.52–0.14––0.073.78
      –––––2.76–0.11––0.363.19
      –––––0.41–0.09––0.551.97
      –––––6.81–0.16––0.522.97
      –––––1.200.020.08––0.482.92
      –––––0.380.020.08––0.702.30
      –––––1.48–0.17––0.303.38
      –––––14.54–0.26–0.340.683.70
      –––––1.670.020.12––0.841.93
      –––––10.780.040.13––0.372.71
      –0.381.810.150.122.520.020.300.120.222.813.89
      –0.143.380.060.073.570.010.220.060.144.431.58
      –0.160.130.100.050.080.010.080.060.070.071.90
      –0.5512.520.250.2514.540.060.920.180.5818.958.19
      212324252728293031323334
      0.41–0.080.190.180.860.113.89–12.40–0.03
      0.080.060.220.040.110.120.072.030.066.540.020.01
      0.12–0.880.060.220.560.302.12–4.78–0.02
      0.240.060.080.110.250.110.083.090.068.310.030.04
      0.410.080.260.030.070.600.042.480.057.560.010.02
      0.290.100.460.050.160.610.225.35–14.82–0.03
      0.10–0.530.060.091.090.212.960.019.15–0.04
      0.29–0.640.050.120.870.222.25–6.80–0.04
      0.59–0.090.070.600.150.161.57–6.24–0.03
      0.17–0.290.050.250.24–1.62–4.07–0.03
      0.13–0.090.060.56–0.041.950.029.69–0.02
      0.25–0.370.140.75–0.022.230.016.78–0.03
      0.80–0.870.040.34–0.731.75–3.41–0.02
      0.80–1.050.110.60–0.761.36–2.69–0.05
      1.08–0.480.040.23–0.751.58–4.550.010.04
      0.57–0.470.040.48–0.641.15–2.65–0.07
      0.190.09–0.040.50–0.012.08–14.210.010.02
      0.590.060.060.081.33–0.011.01–6.55–0.02
      0.520.10–0.071.840.010.011.48–11.57–0.04
      0.670.07–0.041.55–0.010.56–3.79–0.04
      0.350.140.080.081.22–0.011.82–16.27–0.04
      0.140.10–0.071.36–0.011.39–9.24–0.04
      0.570.13–0.080.900.02–1.47–13.630.010.03
      0.400.060.070.071.140.010.011.46–12.40–0.05
      ––––––––––––
      0.390.100.100.090.880.030.021.67–11.29–0.04
      0.090.08–0.071.420.01–0.66–4.89–0.05
      0.300.090.070.081.97––1.39–12.62–0.02
      0.060.080.070.091.16––2.180.0113.87–0.03
      0.350.170.070.102.89–0.011.170.017.56–0.06
      0.090.07–0.091.97––0.78–7.47–0.05
      0.160.080.070.091.85–0.011.220.017.29–0.05
      0.290.100.100.092.390.030.011.060.015.39–0.06
      0.350.100.070.091.66–0.011.140.019.230.020.03
      0.840.090.070.101.79–0.020.92–7.66–0.05
      0.150.070.160.061.45––1.11–6.69–0.04
      0.320.07––1.11––1.17–7.16–0.04
      0.700.100.090.050.770.010.011.24–7.56–0.03
      1.35––0.120.65––1.17–7.12–0.03
      0.400.090.270.080.970.310.161.720.028.260.020.04
      0.300.030.290.030.730.370.240.920.023.570.010.01
      0.060.060.060.030.070.010.010.560.012.650.010.01
      1.350.171.050.192.891.090.765.350.0616.270.030.07
      Totals
      AliphIndoleBenzylMTMSOAOPBenzoxyC3C4C7C8C4PerC8Per
      29.200.410.1918.095.384.301.180.186.584.4517.150.9730.794
      18.790.080.0413.392.931.740.390.915.602.319.010.8600.796
      31.030.120.0614.642.3512.601.100.5319.612.456.800.9740.735
      19.700.240.1113.493.971.310.570.253.073.6011.720.9250.765
      41.230.410.0312.645.5521.610.790.0723.633.3312.110.9970.784
      34.910.310.0522.257.184.001.020.166.136.2720.830.9750.769
      35.990.100.0615.656.8611.491.440.1715.433.7315.140.9890.802
      31.120.290.0514.585.219.561.250.6914.712.7511.360.8550.805
      18.120.620.078.613.215.360.941.164.771.799.230.8040.838
      28.940.190.059.443.0615.920.520.7418.821.966.790.9620.776
      26.210.130.0615.834.854.760.641.017.362.2714.220.8790.862
      31.370.250.1422.776.741.050.811.9813.852.8812.060.8750.807
      22.530.820.0417.732.641.071.090.6013.761.975.310.9580.729
      33.870.860.1124.135.442.891.411.3723.781.915.450.9460.740
      20.181.110.0414.492.871.791.030.929.741.747.040.9140.802
      23.610.580.0417.113.511.801.190.9515.521.465.140.9420.779
      26.520.190.0416.797.102.030.541.040.272.3721.020.2060.899
      28.210.590.0811.154.3411.361.369.620.421.169.840.0420.895
      28.070.520.0714.214.996.971.905.260.191.6716.370.0350.907
      27.260.690.0411.534.1310.001.6012.080.400.676.400.0320.905
      40.850.350.0828.078.173.341.2713.490.732.1622.700.0510.913
      26.340.140.0714.176.903.861.416.600.121.6715.310.0180.902
      28.170.590.0816.148.472.600.962.400.401.7521.820.1430.926
      31.380.400.0720.256.913.011.219.310.381.7418.230.0390.913
      –––––––––––––
      22.460.410.0914.584.342.470.972.531.301.8415.460.3390.894
      20.340.090.079.912.916.041.487.990.220.787.350.0270.904
      25.370.320.0816.744.761.881.995.940.481.5517.220.0750.917
      27.800.060.0917.305.134.171.206.020.412.3818.800.0640.888
      27.650.350.109.563.9211.202.9714.320.081.3111.340.0060.896
      24.930.090.0911.443.497.982.0212.780.360.8910.660.0270.923
      16.860.160.099.622.063.261.925.520.561.319.260.0920.876
      34.550.290.0913.983.9114.162.5023.860.841.228.360.0340.873
      25.420.370.0912.223.008.471.7311.330.491.2212.150.0410.909
      17.300.860.109.822.383.241.865.410.721.009.960.1170.909
      17.630.150.069.813.752.581.495.710.301.2810.070.0500.887
      38.260.320.0023.629.543.951.1524.840.681.4310.860.0270.884
      17.090.720.0511.502.052.720.825.000.861.369.490.1470.875
      24.811.390.1219.443.461.230.6813.280.371.309.830.0270.883
      26.950.410.0715.184.675.731.275.685.602.0212.150.4360.852
      6.570.310.034.671.924.830.566.347.441.104.920.4350.062
      16.860.060.008.612.051.050.390.070.080.675.140.0060.729
      41.231.390.1928.079.5421.612.9724.8423.786.2722.700.9970.926

      Quantities are given in μmol g dry wt−1 and are the mean of three extractions of each ecotype. The nos. at the top of each column refer to the list of glucosinolate in “Materials and Methods.” The values underneath the totals section are as follows: Aliph, sum of aliphatic glucosinolates; Indole, sum of indolic glucosinolates; Benzyl, sum of benzyl glucosinolates; MT, sum of methylthioalkyl glucosinolates; MSO, sum of methylsulfinyl glucosinolates; AOP, sum of alkenyl and hydroxy aliphatic glucosinolates; Benzoxy, sum of benzoyloxy aliphatic glucosinolates; C3, sum of three carbon aliphatic glucosinolates; C4, sum of four carbon aliphatic glucosinolates; C7, sum of seven carbon aliphatic glucosinolates; C8, sum of eight carbon aliphatic glucosinolates; and C4 per, C4/(C3 +C4), C8 per = C8/(C7 + C8).

        • View popup
        Table III.

        Major genetic loci controlling biosynthesis of glucosinolates in Arabidopsis

        EcotypeElongAOPOXOHProfile
        Ag-04323
        Bla-104313
        Bs-14323Allyl
        Cal-04323Butenyl
        Cnt4323HydroxyButenyl
        Ema-14323
        Tac4323
        Pog-04323
        Sorbo4322
        Kas4322Allyl
        Cvi4321Butenyl
        Di-1422–OHPropyl/4MSO
        Aa-0412–
        Col-0412–4MSO-butyl
        Ma-0412–
        Mt-0412–
        Can-0331–
        Kondara332–
        Ei-2332–
        Hodja332–
        Ita-0332–Allyl
        Kil-0332–
        Mr-0332–
        Mrk-0332–
        Su-0331–
        Wi-0332–
        Rsch-0332–
        Bl-1322–
        Di-g322–
        Ka-0322–
        Ler322–
        Lip-0322–OHpropyl
        No-0322–
        Pet322–
        Pi-0322–
        Sei-0322–
        Tsu-1322–
        Yo-0322–
        Oy-0312–3MSOpropyl

        Shown are the genotypes at the five major glucosinolate biosynthetic genetic loci for the 39 ecotypes analyzed in this study. The allele designations are as follows: GS-Elong, 3, C3 producing; 4, C4 producing; GS-AOP, 1, the null methylsulfinylalkyl allele; 2, the hydroxypropyl allele; 3, the alkenyl allele; GS-OX, 1, the Bla-10 allele; 2, the normal wild-type allele; GS-OH, 1, off in all tissues; 2, only on in seeds; and 3, on in all tissues. Ecotypes with no score do not make the necessary precursor for the reaction and are thereby unable to be scored for the allele at that specific locus.

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        Genetic Control of Natural Variation in Arabidopsis Glucosinolate Accumulation
        Daniel J. Kliebenstein, Juergen Kroymann, Paul Brown, Antje Figuth, Deana Pedersen, Jonathan Gershenzon, Thomas Mitchell-Olds
        Plant Physiology Jun 2001, 126 (2) 811-825; DOI: 10.1104/pp.126.2.811

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        Genetic Control of Natural Variation in Arabidopsis Glucosinolate Accumulation
        Daniel J. Kliebenstein, Juergen Kroymann, Paul Brown, Antje Figuth, Deana Pedersen, Jonathan Gershenzon, Thomas Mitchell-Olds
        Plant Physiology Jun 2001, 126 (2) 811-825; DOI: 10.1104/pp.126.2.811
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