CompaRNA - on-line benchmarks of RNA structure prediction methods
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Table of contents:

  1. Overview

  2. Performance Plots

  3. Performance of Multilign(20) - scored higher in this pairwise comparison

  4. Performance of RSpredict(seed) - scored lower in this pairwise comparison

  5. Compile and download dataset for Multilign(20) & RSpredict(seed) [.zip] - may take several seconds...


Overview

Metric Multilign(20) RSpredict(seed)
MCC 0.593 > 0.436
Average MCC ± 95% Confidence Intervals 0.547 ± 0.065 > 0.466 ± 0.058
Sensitivity 0.447 > 0.298
Positive Predictive Value 0.793 > 0.645
Total TP 1367 > 912
Total TN 454838 < 455149
Total FP 543 < 580
Total FP CONTRA 47 < 57
Total FP INCONS 310 < 444
Total FP COMP 186 > 79
Total FN 1692 < 2147
P-value 3.56938820447e-08

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Performance plots


  1. Comparison of performance of Multilign(20) and RSpredict(seed). Positive Predictive Value (PPV) is plotted against sensitivity. Each dot represents a single test of each method. See tables below for raw data (individual counts for Multilign(20) and RSpredict(seed)).

  2. Average Matthews Correlation Coefficients (MCC) with 95% confidence intervals (CIs) were plotted for different RNA families, for which at least 3 members were present in the benchmarking dataset. 'n' denotes the number of MCCs used to calculate the average and CI. See tables below for raw data (individual counts for Multilign(20) and RSpredict(seed)).

  3. Comparison of average Matthews Correlation Coefficients (MCCs) for Multilign(20) and RSpredict(seed). The whiskers correspond to 95% confidence intervals (CIs). 'n' denotes the number of MCCs used to calculate average MCCs and CIs. See tables below for raw data (individual counts for Multilign(20) and RSpredict(seed)).

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Performance of Multilign(20) - scored higher in this pairwise comparison

1. Total counts & total scores for Multilign(20)

Total Base Pair Counts
Total TP 1367
Total TN 454838
Total FP 543
Total FP CONTRA 47
Total FP INCONS 310
Total FP COMP 186
Total FN 1692
Total Scores
MCC 0.593
Average MCC ± 95% Confidence Intervals 0.547 ± 0.065
Sensitivity 0.447
Positive Predictive Value 0.793
Nr of predictions 132

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2. Individual counts for Multilign(20) [ download as .csv ]

RNA Chain Rfam family MCC SENS PPV TP TN FP FP CONTRA FP INCONS FP COMP FN
PDB_00012 1.00 1.00 1.00 7 399 3 0 0 3 0
PDB_00213 0.86 0.78 0.94 32 5016 2 0 2 0 9
PDB_00553 1.00 1.00 1.00 11 454 0 0 0 0 0
PDB_00741 0.87 0.76 1.00 13 690 0 0 0 0 4
PDB_00810 0.67 0.59 0.77 10 1068 3 0 3 0 7
PDB_00828 0.88 0.78 1.00 21 2464 0 0 0 0 6
PDB_00829 0.86 0.75 1.00 18 2260 2 0 0 2 6
PDB_00876 0.97 0.95 1.00 19 971 0 0 0 0 1
PDB_01020 0.86 0.74 1.00 17 2261 3 0 0 3 6
PDB_01050 0.92 0.85 1.00 11 619 2 0 0 2 2
PDB_01073 0.86 0.74 1.00 25 4346 1 0 0 1 9
PDB_01114 0.42 0.37 0.48 10 2829 11 1 10 0 17
PDB_01152 0.88 0.79 1.00 11 550 0 0 0 0 3
PDB_01236 0.88 0.81 0.96 44 11582 3 0 2 1 10
RFA_00389 0.70 0.60 0.82 9 1420 4 0 2 2 6
RFA_00390 1.00 1.00 1.00 15 1416 2 0 0 2 0
RFA_00391 0.57 0.47 0.70 7 1421 4 0 3 1 8
RFA_00396 0.97 0.93 1.00 14 1417 2 0 0 2 1
RFA_00402 1.00 1.00 1.00 15 1416 0 0 0 0 0
RFA_00409 0.70 0.60 0.82 9 1420 4 0 2 2 6
RFA_00416 1.00 1.00 1.00 15 1470 2 0 0 2 0
RFA_00433 1.00 1.00 1.00 15 1416 2 0 0 2 0
RFA_00434 1.00 1.00 1.00 15 1416 2 0 0 2 0
RFA_00436 1.00 1.00 1.00 15 1416 2 0 0 2 0
RFA_00440 1.00 1.00 1.00 15 1470 2 0 0 2 0
RFA_00442 1.00 1.00 1.00 15 1416 2 0 0 2 0
RFA_00444 1.00 1.00 1.00 15 1525 1 0 0 1 0
RFA_00583 0.51 0.53 0.50 8 4934 23 0 8 15 7
RFA_00584 0.87 0.87 0.87 13 2911 14 0 2 12 2
RFA_00585 0.93 0.93 0.93 14 4836 15 0 1 14 1
RFA_00586 0.97 0.93 1.00 14 3902 12 0 0 12 1
RFA_00587 0.93 0.93 0.93 14 4836 15 0 1 14 1
RFA_00588 1.00 1.00 1.00 15 4545 10 0 0 10 0
RFA_00589 1.00 1.00 1.00 15 4356 17 0 0 17 0
RFA_00594 0.83 0.80 0.86 12 2761 14 0 2 12 3
RFA_00596 0.74 0.67 0.83 10 5983 18 0 2 16 5
RFA_00658 0.00 0.00 0.00 0 1128 0 0 0 0 14
RFA_00664 0.00 0.00 0.00 0 990 0 0 0 0 14
RFA_00667 0.00 0.00 0.00 0 990 0 0 0 0 14
RFA_00668 0.00 0.00 0.00 0 990 0 0 0 0 14
RFA_00672 0.00 0.00 0.00 0 903 0 0 0 0 13
RFA_00673 0.00 0.00 0.00 0 1128 0 0 0 0 14
RFA_00674 0.00 0.00 0.00 0 1128 0 0 0 0 14
RFA_00675 0.00 0.00 0.00 0 990 0 0 0 0 14
RFA_00677 0.00 0.00 0.00 0 990 0 0 0 0 14
RFA_00678 0.00 0.00 0.00 0 946 0 0 0 0 14
RFA_00680 0.00 0.00 0.00 0 1128 0 0 0 0 14
RFA_00704 0.00 0.00 0.00 0 990 0 0 0 0 14
RFA_00707 0.00 0.00 0.00 0 1035 0 0 0 0 14
RFA_00715 0.00 0.00 0.00 0 946 0 0 0 0 14
RFA_00717 0.00 0.00 0.00 0 903 0 0 0 0 14
RFA_00730 0.69 0.67 0.73 8 892 4 1 2 1 4
RFA_00731 1.00 1.00 1.00 12 891 1 0 0 1 0
RFA_00733 1.00 1.00 1.00 12 891 1 0 0 1 0
RFA_00734 0.78 0.75 0.82 9 892 3 0 2 1 3
RFA_00736 0.69 0.67 0.73 8 892 4 1 2 1 4
RFA_00737 0.73 0.67 0.80 8 893 3 0 2 1 4
RFA_00745 0.69 0.67 0.73 8 935 4 1 2 1 4
RFA_00749 0.69 0.67 0.73 8 892 4 1 2 1 4
RFA_00758 0.63 0.58 0.70 7 893 3 1 2 0 5
RFA_00762 0.69 0.67 0.73 8 892 3 0 3 0 4
RFA_00763 1.00 1.00 1.00 12 891 0 0 0 0 0
RFA_00764 0.69 0.67 0.73 8 892 4 1 2 1 4
RFA_00765 0.91 0.83 1.00 10 893 1 0 0 1 2
RFA_00786 0.64 0.66 0.64 21 5017 12 2 10 0 11
RFA_00791 0.89 0.84 0.93 27 5122 3 0 2 1 5
RFA_00792 0.94 0.94 0.94 30 5018 4 0 2 2 2
RFA_00801 0.81 0.78 0.83 25 5020 6 0 5 1 7
SPR_00273 0.76 0.76 0.76 16 2829 5 0 5 0 5
SPR_00277 0.98 0.95 1.00 20 2906 0 0 0 0 1
SPR_00721 1.00 1.00 1.00 21 2905 0 0 0 0 0
SPR_01157 0.83 0.81 0.85 17 2755 4 0 3 1 4
SRP_00020 0.29 0.15 0.56 5 5556 4 0 4 0 28
SRP_00038 0.89 0.85 0.93 28 5020 2 0 2 0 5
SRP_00042 0.36 0.24 0.55 6 3994 6 1 4 1 19
SRP_00046 0.00 0.00 0.00 0 4753 0 0 0 0 32
SRP_00051 0.00 0.00 0.00 0 5450 10 0 10 0 30
SRP_00054 0.47 0.22 1.00 7 5558 0 0 0 0 25
SRP_00057 0.34 0.20 0.58 7 6316 5 1 4 0 28
SRP_00058 0.73 0.68 0.79 23 5749 6 1 5 0 11
SRP_00072 0.00 0.00 0.00 0 6318 10 0 10 0 29
SRP_00074 0.00 0.00 0.00 0 6543 12 0 12 0 29
SRP_00077 0.57 0.32 1.00 12 6658 0 0 0 0 25
SRP_00081 0.00 0.00 0.00 0 5555 10 2 8 0 31
SRP_00084 0.36 0.21 0.64 7 5449 4 0 4 0 27
SRP_00094 0.73 0.75 0.72 18 4070 7 1 6 0 6
SRP_00095 0.30 0.28 0.33 7 3984 14 4 10 0 18
SRP_00098 0.00 0.00 0.00 0 6430 11 1 10 0 37
SRP_00107 0.27 0.32 0.24 6 3630 21 4 15 2 13
SRP_00121 0.60 0.40 0.91 10 4267 1 0 1 0 15
SRP_00128 0.14 0.08 0.25 3 6774 9 0 9 0 33
SRP_00132 0.28 0.17 0.45 5 4840 7 0 6 1 24
SRP_00134 0.51 0.55 0.48 16 5962 23 3 14 6 13
SRP_00141 0.65 0.51 0.84 21 6303 4 1 3 0 20
SRP_00161 0.71 0.50 1.00 11 2839 0 0 0 0 11
SRP_00162 0.87 0.87 0.87 20 2903 4 0 3 1 3
SRP_00163 0.52 0.27 1.00 7 3479 0 0 0 0 19
SRP_00164 0.00 0.00 0.00 0 6892 11 2 9 0 35
SRP_00170 0.70 0.56 0.88 14 4079 3 1 1 1 11
SRP_00194 0.00 0.00 0.00 0 3240 0 0 0 0 21
SRP_00209 0.20 0.19 0.23 5 4634 17 4 13 0 22
SRP_00213 0.54 0.36 0.82 9 4175 2 0 2 0 16
SRP_00215 0.53 0.32 0.89 8 3231 1 0 1 0 17
SRP_00227 0.94 0.94 0.94 29 5120 2 1 1 0 2
SRP_00231 0.46 0.26 0.82 9 5449 2 0 2 0 25
SRP_00243 0.47 0.23 1.00 7 5144 0 0 0 0 24
SRP_00244 0.00 0.00 0.00 0 6892 11 1 10 0 39
SRP_00250 0.00 0.00 0.00 0 6786 0 0 0 0 36
SRP_00251 0.77 0.67 0.89 24 6994 3 1 2 0 12
SRP_00266 0.00 0.00 0.00 0 4656 0 0 0 0 29
SRP_00268 0.93 0.93 0.93 25 4159 2 1 1 0 2
SRP_00269 0.00 0.00 0.00 0 5043 7 1 6 0 33
SRP_00270 0.80 0.65 1.00 20 4636 0 0 0 0 11
SRP_00271 0.00 0.00 0.00 0 3240 0 0 0 0 24
SRP_00273 0.55 0.30 1.00 12 6774 0 0 0 0 28
SRP_00274 0.95 0.95 0.95 36 6865 2 1 1 0 2
SRP_00303 0.31 0.09 1.00 3 6667 0 0 0 0 29
SRP_00310 0.84 0.83 0.86 19 2904 4 0 3 1 4
SRP_00312 0.77 0.67 0.90 18 4166 2 1 1 0 9
SRP_00318 0.62 0.39 1.00 9 2994 0 0 0 0 14
SRP_00325 0.70 0.58 0.86 19 4349 3 0 3 0 14
SRP_00333 0.00 0.00 0.00 0 5151 0 0 0 0 34
SRP_00338 0.00 0.00 0.00 0 5460 0 0 0 0 36
SRP_00341 0.00 0.00 0.00 0 4741 12 3 9 0 31
SRP_00342 0.46 0.28 0.78 7 3996 4 0 2 2 18
SRP_00350 0.39 0.26 0.58 7 3904 5 0 5 0 20
SRP_00356 0.58 0.37 0.92 11 5038 1 0 1 0 19
SRP_00357 0.00 0.00 0.00 0 5356 0 0 0 0 36
SRP_00358 0.32 0.14 0.71 5 6321 2 1 1 0 30
SRP_00367 0.00 0.00 0.00 0 6775 11 1 10 0 35
SRP_00369 0.58 0.34 1.00 12 5766 0 0 0 0 23
SRP_00383 0.54 0.29 1.00 7 3233 0 0 0 0 17

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Performance of RSpredict(seed) - scored lower in this pairwise comparison

1. Total counts & total scores for RSpredict(seed)

Total Base Pair Counts
Total TP 912
Total TN 455149
Total FP 580
Total FP CONTRA 57
Total FP INCONS 444
Total FP COMP 79
Total FN 2147
Total Scores
MCC 0.436
Average MCC ± 95% Confidence Intervals 0.466 ± 0.058
Sensitivity 0.298
Positive Predictive Value 0.645
Nr of predictions 132

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2. Individual counts for RSpredict(seed) [ download as .csv ]

RNA Chain Rfam family MCC SENS PPV TP TN FP FP CONTRA FP INCONS FP COMP FN
PDB_00012 1.00 1.00 1.00 7 399 1 0 0 1 0
PDB_00213 0.56 0.49 0.65 20 5019 11 0 11 0 21
PDB_00553 0.90 0.82 1.00 9 456 0 0 0 0 2
PDB_00741 0.00 0.00 0.00 0 703 0 0 0 0 17
PDB_00810 0.80 0.65 1.00 11 1070 0 0 0 0 6
PDB_00828 0.54 0.41 0.73 11 2470 4 2 2 0 16
PDB_00829 0.74 0.63 0.88 15 2261 2 2 0 0 9
PDB_00876 0.57 0.50 0.67 10 975 5 0 5 0 10
PDB_01020 0.76 0.65 0.88 15 2261 2 2 0 0 8
PDB_01050 0.62 0.46 0.86 6 623 2 0 1 1 7
PDB_01073 0.39 0.18 0.86 6 4364 1 0 1 0 28
PDB_01114 0.74 0.56 1.00 15 2835 0 0 0 0 12
PDB_01152 0.88 0.79 1.00 11 550 0 0 0 0 3
PDB_01236 0.00 0.00 0.00 0 11621 7 0 7 0 54
RFA_00389 0.63 0.40 1.00 6 1425 1 0 0 1 9
RFA_00390 0.63 0.40 1.00 6 1425 1 0 0 1 9
RFA_00391 0.63 0.40 1.00 6 1425 1 0 0 1 9
RFA_00396 0.63 0.40 1.00 6 1425 1 0 0 1 9
RFA_00402 0.63 0.40 1.00 6 1425 0 0 0 0 9
RFA_00409 0.58 0.33 1.00 5 1426 1 0 0 1 10
RFA_00416 0.63 0.40 1.00 6 1479 1 0 0 1 9
RFA_00433 0.63 0.40 1.00 6 1425 1 0 0 1 9
RFA_00434 0.63 0.40 1.00 6 1425 1 0 0 1 9
RFA_00436 0.63 0.40 1.00 6 1425 1 0 0 1 9
RFA_00440 0.63 0.40 1.00 6 1479 1 0 0 1 9
RFA_00442 0.63 0.40 1.00 6 1425 1 0 0 1 9
RFA_00444 0.00 0.00 0.00 0 1538 2 0 2 0 15
RFA_00583 1.00 1.00 1.00 15 4935 7 0 0 7 0
RFA_00584 1.00 1.00 1.00 15 2911 2 0 0 2 0
RFA_00585 1.00 1.00 1.00 15 4836 6 0 0 6 0
RFA_00586 0.97 0.93 1.00 14 3902 2 0 0 2 1
RFA_00587 1.00 1.00 1.00 15 4836 4 0 0 4 0
RFA_00588 0.97 0.93 1.00 14 4546 4 0 0 4 1
RFA_00589 0.93 0.87 1.00 13 4358 3 0 0 3 2
RFA_00594 0.93 0.87 1.00 13 2762 2 0 0 2 2
RFA_00596 0.82 0.67 1.00 10 5985 5 0 0 5 5
RFA_00658 0.64 0.57 0.73 8 1117 4 0 3 1 6
RFA_00664 0.59 0.50 0.70 7 980 4 0 3 1 7
RFA_00667 0.64 0.57 0.73 8 979 4 0 3 1 6
RFA_00668 0.71 0.57 0.89 8 981 1 0 1 0 6
RFA_00672 0.61 0.54 0.70 7 893 3 0 3 0 6
RFA_00673 0.42 0.36 0.50 5 1118 6 0 5 1 9
RFA_00674 0.69 0.64 0.75 9 1116 4 0 3 1 5
RFA_00675 0.62 0.50 0.78 7 981 3 0 2 1 7
RFA_00677 0.72 0.64 0.82 9 979 2 0 2 0 5
RFA_00678 0.43 0.29 0.67 4 940 3 0 2 1 10
RFA_00680 0.69 0.64 0.75 9 1116 4 0 3 1 5
RFA_00704 0.35 0.29 0.44 4 981 6 0 5 1 10
RFA_00707 0.67 0.57 0.80 8 1025 3 0 2 1 6
RFA_00715 0.66 0.50 0.88 7 938 2 0 1 1 7
RFA_00717 0.50 0.36 0.71 5 896 2 0 2 0 9
RFA_00730 0.91 0.83 1.00 10 893 1 0 0 1 2
RFA_00731 0.91 0.83 1.00 10 893 1 0 0 1 2
RFA_00733 0.91 0.83 1.00 10 893 1 0 0 1 2
RFA_00734 0.91 0.83 1.00 10 893 0 0 0 0 2
RFA_00736 0.91 0.83 1.00 10 893 1 0 0 1 2
RFA_00737 0.91 0.83 1.00 10 893 1 0 0 1 2
RFA_00745 0.91 0.83 1.00 10 936 1 0 0 1 2
RFA_00749 0.91 0.83 1.00 10 893 1 0 0 1 2
RFA_00758 0.86 0.75 1.00 9 894 0 0 0 0 3
RFA_00762 0.91 0.83 1.00 10 893 0 0 0 0 2
RFA_00763 0.91 0.83 1.00 10 893 1 0 0 1 2
RFA_00764 0.91 0.83 1.00 10 893 1 0 0 1 2
RFA_00765 0.91 0.83 1.00 10 893 1 0 0 1 2
RFA_00786 0.61 0.59 0.63 19 5020 12 2 9 1 13
RFA_00791 0.58 0.53 0.63 17 5124 11 2 8 1 15
RFA_00792 0.64 0.63 0.67 20 5020 11 2 8 1 12
RFA_00801 0.59 0.56 0.62 18 5021 12 2 9 1 14
SPR_00273 0.00 0.00 0.00 0 2846 4 0 4 0 21
SPR_00277 0.00 0.00 0.00 0 2923 4 0 3 1 21
SPR_00721 0.00 0.00 0.00 0 2923 3 0 3 0 21
SPR_01157 0.00 0.00 0.00 0 2774 1 0 1 0 21
SRP_00020 0.38 0.24 0.62 8 5552 5 0 5 0 25
SRP_00038 0.35 0.21 0.58 7 5038 5 1 4 0 26
SRP_00042 0.00 0.00 0.00 0 3997 8 0 8 0 25
SRP_00046 0.33 0.19 0.60 6 4743 4 1 3 0 26
SRP_00051 0.00 0.00 0.00 0 5448 12 1 11 0 30
SRP_00054 0.00 0.00 0.00 0 5555 10 1 9 0 32
SRP_00057 0.21 0.11 0.40 4 6318 6 0 6 0 31
SRP_00058 0.31 0.18 0.55 6 5767 6 0 5 1 28
SRP_00072 0.00 0.00 0.00 0 6320 8 4 4 0 29
SRP_00074 0.00 0.00 0.00 0 6548 7 1 6 0 29
SRP_00077 0.00 0.00 0.00 0 6660 10 1 9 0 37
SRP_00081 0.29 0.19 0.43 6 5551 8 1 7 0 25
SRP_00084 0.34 0.21 0.58 7 5448 5 1 4 0 27
SRP_00094 0.00 0.00 0.00 0 4089 6 1 5 0 24
SRP_00095 0.57 0.44 0.73 11 3990 4 3 1 0 14
SRP_00098 0.04 0.03 0.08 1 6429 11 0 11 0 36
SRP_00107 0.43 0.32 0.60 6 3645 4 0 4 0 13
SRP_00121 0.60 0.36 1.00 9 4269 0 0 0 0 16
SRP_00128 0.00 0.00 0.00 0 6778 8 2 6 0 36
SRP_00132 0.32 0.21 0.50 6 4839 6 2 4 0 23
SRP_00134 0.65 0.66 0.66 19 5966 11 0 10 1 10
SRP_00141 0.26 0.15 0.46 6 6315 7 0 7 0 35
SRP_00161 0.41 0.32 0.54 7 2837 6 0 6 0 15
SRP_00162 0.00 0.00 0.00 0 2921 5 4 1 0 23
SRP_00163 0.43 0.27 0.70 7 3476 3 0 3 0 19
SRP_00164 0.72 0.60 0.88 21 6879 4 1 2 1 14
SRP_00170 0.00 0.00 0.00 0 4091 4 0 4 0 25
SRP_00194 0.16 0.10 0.29 2 3233 5 2 3 0 19
SRP_00209 0.00 0.00 0.00 0 4649 7 1 6 0 27
SRP_00213 0.39 0.28 0.54 7 4173 6 1 5 0 18
SRP_00215 0.44 0.28 0.70 7 3230 3 1 2 0 18
SRP_00227 0.00 0.00 0.00 0 5143 8 0 8 0 31
SRP_00231 0.46 0.29 0.71 10 5446 4 0 4 0 24
SRP_00243 0.00 0.00 0.00 0 5140 11 1 10 0 31
SRP_00244 0.00 0.00 0.00 0 6891 12 1 11 0 39
SRP_00250 0.00 0.00 0.00 0 6775 11 2 9 0 36
SRP_00251 0.00 0.00 0.00 0 7011 10 1 9 0 36
SRP_00266 0.00 0.00 0.00 0 4647 9 1 8 0 29
SRP_00268 0.00 0.00 0.00 0 4180 6 1 5 0 27
SRP_00269 0.29 0.18 0.46 6 5037 7 0 7 0 27
SRP_00270 0.28 0.16 0.50 5 4646 5 0 5 0 26
SRP_00271 0.00 0.00 0.00 0 3235 5 0 5 0 24
SRP_00273 0.00 0.00 0.00 0 6778 8 1 7 0 40
SRP_00274 0.00 0.00 0.00 0 6892 11 1 10 0 38
SRP_00303 0.00 0.00 0.00 0 6665 5 1 4 0 32
SRP_00310 0.60 0.43 0.83 10 2914 3 0 2 1 13
SRP_00312 0.00 0.00 0.00 0 4178 8 0 8 0 27
SRP_00318 0.69 0.48 1.00 11 2992 0 0 0 0 12
SRP_00325 0.36 0.21 0.64 7 4360 4 0 4 0 26
SRP_00333 0.33 0.21 0.54 7 5138 6 0 6 0 27
SRP_00338 0.51 0.33 0.80 12 5445 4 0 3 1 24
SRP_00341 0.35 0.23 0.54 7 4740 6 1 5 0 24
SRP_00342 0.38 0.24 0.60 6 3995 4 0 4 0 19
SRP_00350 0.19 0.07 0.50 2 3912 2 0 2 0 25
SRP_00356 0.00 0.00 0.00 0 5043 7 1 6 0 30
SRP_00357 0.46 0.31 0.69 11 5340 5 0 5 0 25
SRP_00358 0.49 0.29 0.83 10 6316 2 1 1 0 25
SRP_00367 0.00 0.00 0.00 0 6774 12 0 12 0 35
SRP_00369 0.30 0.17 0.55 6 5767 5 0 5 0 29
SRP_00383 0.54 0.42 0.71 10 3226 4 0 4 0 14

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Matthews Correlation Coeffient, Sensitivity and Positive Predictive Value have been calculated based on the paper by Gardener & Giegerich, 2004.