==== Secondary Structure Definition by the program DSSP, updated CMBI version by ElmK / April 1,2000 ==== DATE=6-JAN-2010 . REFERENCE W. KABSCH AND C.SANDER, BIOPOLYMERS 22 (1983) 2577-2637 . HEADER HYDROLASE 02-APR-09 2WF1 . COMPND 2 MOLECULE: BETA-SECRETASE 1; . SOURCE 2 ORGANISM_SCIENTIFIC: HOMO SAPIENS; . AUTHOR N.CHARRIER,B.CLARKE,E.DEMONT,C.DINGWALL,R.DUNSDON,J.HAWKINS, . 374 2 3 2 1 TOTAL NUMBER OF RESIDUES, NUMBER OF CHAINS, NUMBER OF SS-BRIDGES(TOTAL,INTRACHAIN,INTERCHAIN) . 16189.0 ACCESSIBLE SURFACE OF PROTEIN (ANGSTROM**2) . 257 68.7 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(J) , SAME NUMBER PER 100 RESIDUES . 18 4.8 TOTAL NUMBER OF HYDROGEN BONDS IN PARALLEL BRIDGES, SAME NUMBER PER 100 RESIDUES . 116 31.0 TOTAL NUMBER OF HYDROGEN BONDS IN ANTIPARALLEL BRIDGES, SAME NUMBER PER 100 RESIDUES . 3 0.8 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I-5), SAME NUMBER PER 100 RESIDUES . 2 0.5 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I-4), SAME NUMBER PER 100 RESIDUES . 3 0.8 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I-3), SAME NUMBER PER 100 RESIDUES . 0 0.0 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I-2), SAME NUMBER PER 100 RESIDUES . 1 0.3 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I-1), SAME NUMBER PER 100 RESIDUES . 0 0.0 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I+0), SAME NUMBER PER 100 RESIDUES . 0 0.0 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I+1), SAME NUMBER PER 100 RESIDUES . 45 12.0 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I+2), SAME NUMBER PER 100 RESIDUES . 38 10.2 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I+3), SAME NUMBER PER 100 RESIDUES . 24 6.4 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I+4), SAME NUMBER PER 100 RESIDUES . 4 1.1 TOTAL NUMBER OF HYDROGEN BONDS OF TYPE O(I)-->H-N(I+5), SAME NUMBER PER 100 RESIDUES . 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 *** HISTOGRAMS OF *** . 0 0 0 3 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 RESIDUES PER ALPHA HELIX . 0 0 2 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 PARALLEL BRIDGES PER LADDER . 2 4 3 3 2 5 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ANTIPARALLEL BRIDGES PER LADDER . 0 1 0 2 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 LADDERS PER SHEET . # RESIDUE AA STRUCTURE BP1 BP2 ACC N-H-->O O-->H-N N-H-->O O-->H-N TCO KAPPA ALPHA PHI PSI X-CA Y-CA Z-CA 1 62 A E 0 0 175 0, 0.0 0, 0.0 0, 0.0 0, 0.0 0.000 360.0 360.0 360.0 -27.8 3.4 -6.1 25.7 2 63 A M > + 0 0 24 164,-0.0 3,-1.1 161,-0.0 162,-0.6 0.570 360.0 115.1 -90.1 -11.4 6.6 -4.4 27.0 3 64 A V T 3 S+ 0 0 67 1,-0.2 160,-0.1 160,-0.1 162,-0.0 -0.346 82.2 12.0 -62.8 138.4 6.7 -6.5 30.2 4 65 A D T 3 S+ 0 0 80 149,-0.1 -1,-0.2 1,-0.1 159,-0.2 0.806 82.8 134.2 64.7 33.0 9.7 -8.8 30.6 5 66 A N < + 0 0 17 -3,-1.1 12,-0.5 84,-0.1 2,-0.4 0.373 51.4 80.9 -93.2 4.1 11.5 -7.2 27.6 6 67 A L E +AB 16 162A 0 156,-2.5 156,-1.5 10,-0.1 2,-0.3 -0.877 52.4 171.2-107.9 144.9 14.8 -7.0 29.5 7 68 A R E +AB 15 161A 103 8,-2.1 8,-3.1 -2,-0.4 2,-0.3 -0.878 9.2 142.1-134.2 172.4 17.4 -9.8 30.0 8 69 A G E -AB 14 160A 4 152,-1.5 152,-3.1 -2,-0.3 2,-0.4 -0.970 43.7 -99.0 175.0 176.9 20.8 -10.0 31.4 9 70 A K E > - B 0 159A 75 4,-0.7 3,-2.4 -2,-0.3 4,-0.2 -0.964 54.8 -89.8-117.1 141.8 23.5 -11.8 33.5 10 71 A S T 3 S+ 0 0 21 148,-2.1 290,-0.1 -2,-0.4 146,-0.0 -0.173 112.0 9.6 -51.9 132.0 24.1 -10.9 37.1 11 72 A G T 3 S+ 0 0 26 211,-0.1 -1,-0.3 1,-0.1 211,-0.2 0.496 114.9 82.3 80.9 2.9 26.8 -8.2 37.4 12 73 A Q S < S- 0 0 54 -3,-2.4 -2,-0.2 1,-0.3 98,-0.1 0.224 95.2-104.3-128.0 13.1 26.8 -7.6 33.6 13 74 A G - 0 0 11 -4,-0.2 -4,-0.7 -5,-0.1 2,-0.4 -0.105 30.5 -84.0 94.0 169.3 23.9 -5.2 33.0 14 75 A Y E -A 8 0A 9 17,-0.4 17,-2.8 205,-0.3 2,-0.3 -0.956 42.5-164.9-116.5 138.4 20.4 -5.5 31.6 15 76 A Y E -AC 7 30A 27 -8,-3.1 -8,-2.1 -2,-0.4 2,-0.3 -0.908 11.6-154.9-128.6 155.4 19.7 -5.3 27.9 16 77 A V E -AC 6 29A 0 13,-2.5 13,-2.9 -2,-0.3 2,-0.3 -0.878 31.2-106.9-120.8 151.2 16.7 -4.8 25.5 17 78 A E E + C 0 28A 66 -12,-0.5 72,-0.4 -2,-0.3 2,-0.3 -0.649 45.7 168.1 -80.8 134.9 16.4 -5.9 21.9 18 79 A M E - C 0 27A 0 9,-2.8 9,-2.8 -2,-0.3 2,-0.4 -0.923 25.3-139.6-135.9 163.9 16.7 -3.2 19.3 19 80 A T E -DC 86 26A 13 67,-2.3 67,-2.0 -2,-0.3 2,-0.4 -0.979 14.4-168.0-123.1 144.9 17.1 -3.0 15.6 20 81 A V E > +DC 85 25A 1 5,-2.2 5,-2.0 -2,-0.4 65,-0.2 -0.988 46.4 4.9-128.2 141.2 19.3 -0.5 13.7 21 82 A G T 5S- 0 0 4 63,-2.6 4,-0.1 -2,-0.4 36,-0.1 -0.056 88.4 -48.2 88.4 174.2 19.4 0.3 10.0 22 83 A S T 5S+ 0 0 32 2,-0.1 36,-2.2 1,-0.1 37,-0.4 -0.977 129.5 33.2-133.5 118.8 17.4 -0.7 6.9 23 84 A P T 5S- 0 0 96 0, 0.0 -1,-0.1 0, 0.0 -3,-0.1 0.578 120.7-102.0 -64.7 158.3 16.9 -3.6 6.6 24 85 A P T 5 - 0 0 91 0, 0.0 2,-0.5 0, 0.0 -3,-0.2 -0.276 31.6-158.1 -61.0 129.4 16.7 -4.0 10.4 25 86 A Q E < -C 20 0A 29 -5,-2.0 -5,-2.2 -4,-0.1 2,-0.3 -0.948 17.1-135.7-106.2 123.1 19.8 -5.4 12.1 26 87 A T E +C 19 0A 53 -2,-0.5 2,-0.3 -7,-0.2 -7,-0.2 -0.615 31.5 165.4 -84.7 132.4 19.0 -6.9 15.5 27 88 A L E -C 18 0A 3 -9,-2.8 -9,-2.8 -2,-0.3 2,-0.5 -0.983 36.7-123.3-140.0 150.6 21.2 -6.2 18.5 28 89 A N E -C 17 0A 23 -2,-0.3 88,-3.1 -11,-0.2 89,-0.9 -0.864 33.2-162.6 -93.0 129.6 21.0 -6.5 22.2 29 90 A I E -Ce 16 117A 0 -13,-2.9 -13,-2.5 -2,-0.5 89,-0.3 -0.950 16.6-127.1-124.4 123.8 21.6 -3.2 23.9 30 91 A L E -Ce 15 118A 10 87,-3.0 89,-3.4 -2,-0.5 2,-0.6 -0.465 24.6-129.6 -68.6 132.7 22.4 -2.5 27.5 31 92 A V E + e 0 119A 2 -17,-2.8 2,-0.5 -2,-0.2 -17,-0.4 -0.750 31.6 179.9 -86.9 117.4 20.1 -0.0 29.2 32 93 A D E > + e 0 120A 7 87,-2.5 89,-2.8 -2,-0.6 3,-1.0 -0.885 25.2 176.2-129.2 102.8 22.1 2.7 30.9 33 94 A T T 3 S+ 0 0 0 -2,-0.5 156,-0.5 1,-0.2 184,-0.2 0.492 80.6 72.6 -77.2 -0.4 20.6 5.6 32.8 34 95 A G T 3 S+ 0 0 10 154,-0.1 2,-0.3 88,-0.1 -1,-0.2 0.307 101.5 30.4 -99.2 8.2 24.1 6.6 33.7 35 96 A S < - 0 0 11 -3,-1.0 87,-2.7 84,-0.2 92,-0.1 -0.919 62.6-138.5-153.5-179.6 25.2 7.9 30.3 36 97 A S + 0 0 2 90,-0.4 2,-0.4 -2,-0.3 100,-0.2 0.288 63.6 97.9-135.6 10.3 23.8 9.6 27.2 37 98 A N - 0 0 6 89,-0.1 83,-1.0 98,-0.1 2,-0.6 -0.878 56.6-146.3-104.6 135.6 25.3 8.0 24.1 38 99 A F E +F 119 0A 8 -2,-0.4 62,-2.5 81,-0.2 2,-0.3 -0.893 35.6 164.6 -92.2 119.5 23.7 5.3 22.0 39 100 A A E -Fg 118 100A 0 79,-2.4 79,-2.9 -2,-0.6 2,-0.3 -0.992 17.8-170.5-141.6 146.0 26.6 3.2 20.7 40 101 A V E -Fg 117 101A 0 60,-2.5 62,-2.9 -2,-0.3 77,-0.3 -0.988 36.9 -96.4-138.2 144.8 26.9 -0.2 19.1 41 102 A G E + g 0 102A 1 75,-3.3 11,-3.2 -2,-0.3 62,-0.1 -0.393 42.0 172.6 -54.3 134.1 29.7 -2.6 18.2 42 103 A A + 0 0 9 60,-2.3 12,-0.2 9,-0.3 -1,-0.1 0.123 57.7 45.8-138.1 16.3 30.4 -2.0 14.6 43 104 A A S S- 0 0 27 59,-0.2 -1,-0.1 7,-0.1 9,-0.1 -0.970 98.7 -74.8-154.6 157.1 33.5 -4.1 14.0 44 105 A P + 0 0 100 0, 0.0 -2,-0.1 0, 0.0 5,-0.1 -0.211 50.2 173.0 -58.4 142.4 34.6 -7.6 15.0 45 106 A H > - 0 0 55 3,-0.3 3,-1.9 1,-0.1 59,-0.0 -0.985 41.9-118.9-148.9 145.2 35.5 -8.2 18.6 46 107 A P T 3 S+ 0 0 116 0, 0.0 -1,-0.1 0, 0.0 0, 0.0 0.826 116.0 43.6 -55.3 -31.9 36.3 -11.4 20.5 47 108 A F T 3 S+ 0 0 74 64,-0.1 2,-0.6 -3,-0.0 65,-0.3 0.367 94.6 95.3 -97.5 4.0 33.4 -10.9 22.8 48 109 A L < - 0 0 19 -3,-1.9 -3,-0.3 1,-0.1 3,-0.1 -0.864 48.4-171.4-100.8 123.9 30.8 -9.8 20.2 49 110 A H S S- 0 0 131 -2,-0.6 2,-0.3 1,-0.2 -1,-0.1 0.751 79.8 -4.5 -77.4 -26.2 28.5 -12.4 18.7 50 111 A R - 0 0 89 -7,-0.1 2,-0.3 2,-0.0 -1,-0.2 -0.961 67.6-160.7-157.5 166.8 27.4 -9.9 16.1 51 112 A Y - 0 0 77 -2,-0.3 2,-0.3 -9,-0.1 -9,-0.3 -0.980 33.8 -82.9-154.4 165.4 27.8 -6.2 15.2 52 113 A Y - 0 0 2 -11,-3.2 2,-0.7 -2,-0.3 -31,-0.1 -0.541 35.0-159.7 -72.1 123.8 26.3 -3.3 13.3 53 114 A Q > - 0 0 65 -2,-0.3 3,-1.5 1,-0.2 4,-0.2 -0.864 7.3-173.7-109.4 99.5 27.3 -3.4 9.6 54 115 A R G > S+ 0 0 57 -2,-0.7 3,-1.7 1,-0.3 6,-0.3 0.800 79.2 65.6 -63.0 -31.1 26.8 0.1 8.2 55 116 A Q G 3 S+ 0 0 140 1,-0.3 -1,-0.3 3,-0.0 -2,-0.0 0.726 99.0 54.1 -66.5 -21.4 27.5 -0.9 4.6 56 117 A L G < S+ 0 0 97 -3,-1.5 2,-0.6 1,-0.0 -1,-0.3 0.431 92.7 85.2 -92.1 -0.7 24.3 -3.0 4.7 57 118 A S X - 0 0 9 -3,-1.7 3,-0.7 -4,-0.2 -36,-0.1 -0.904 53.5-170.4-108.7 118.4 22.1 -0.1 5.8 58 119 A S T 3 S+ 0 0 108 -36,-2.2 -1,-0.1 -2,-0.6 -4,-0.0 0.689 89.3 55.6 -75.9 -18.1 20.7 2.2 3.0 59 120 A T T 3 S+ 0 0 30 -37,-0.4 -1,-0.2 25,-0.1 2,-0.2 0.469 80.3 115.9 -96.2 -3.4 19.5 4.6 5.7 60 121 A Y < - 0 0 57 -3,-0.7 2,-0.4 -6,-0.3 23,-0.2 -0.438 44.8-168.2 -66.7 135.3 22.8 5.1 7.4 61 122 A R E -H 82 0A 151 21,-2.6 21,-2.4 -2,-0.2 2,-0.4 -0.991 15.4-137.7-127.8 125.6 24.3 8.6 7.3 62 123 A D E -H 81 0A 61 -2,-0.4 19,-0.3 19,-0.2 18,-0.0 -0.643 7.0-163.2 -79.4 130.1 27.9 9.3 8.3 63 124 A L E - 0 0 59 17,-2.2 18,-0.2 -2,-0.4 -1,-0.1 0.434 33.9-130.9 -91.2 -0.3 28.3 12.4 10.4 64 125 A R E + 0 0 215 16,-0.5 2,-0.3 1,-0.2 17,-0.1 0.913 68.5 110.4 52.7 51.5 32.0 12.4 9.6 65 126 A K E - 0 0 120 15,-0.5 15,-3.2 2,-0.0 -2,-0.2 -0.960 57.7-138.3-155.1 133.9 33.1 12.9 13.2 66 127 A G E -H 79 0A 41 -2,-0.3 2,-0.3 13,-0.2 13,-0.2 -0.513 17.0-162.7 -90.8 163.4 34.8 10.6 15.7 67 128 A V E -H 78 0A 12 11,-2.3 11,-2.6 -2,-0.2 2,-0.4 -0.998 8.9-160.1-148.6 144.3 33.9 10.2 19.3 68 129 A Y E -H 77 0A 113 -2,-0.3 9,-0.2 9,-0.2 -2,-0.0 -0.991 8.9-174.0-127.3 132.2 35.6 8.8 22.5 69 130 A V E -H 76 0A 8 7,-2.7 7,-1.9 -2,-0.4 2,-0.5 -0.923 3.7-170.3-135.7 108.3 33.7 7.7 25.6 70 131 A P E -H 75 0A 45 0, 0.0 5,-0.2 0, 0.0 2,-0.2 -0.844 10.0-172.1 -91.5 128.0 35.4 6.6 28.9 71 132 A Y - 0 0 53 3,-2.9 3,-0.2 -2,-0.5 -36,-0.0 -0.511 45.1 -94.8-103.4-179.6 33.2 5.1 31.6 72 133 A T S S+ 0 0 97 1,-0.3 2,-0.5 -2,-0.2 -1,-0.1 0.935 123.7 28.2 -58.8 -46.3 34.1 4.2 35.1 73 134 A Q S S- 0 0 142 1,-0.0 -1,-0.3 35,-0.0 34,-0.0 -0.981 126.7 -8.7-115.2 122.2 34.6 0.7 33.7 74 135 A G - 0 0 16 -2,-0.5 -3,-2.9 -3,-0.2 2,-0.3 0.159 59.0-140.7 86.7 158.9 35.7 0.4 30.1 75 136 A K E -HI 70 106A 86 31,-1.6 31,-2.8 -5,-0.2 2,-0.3 -0.983 12.0-171.1-152.4 157.6 36.0 2.8 27.2 76 137 A W E -HI 69 105A 3 -7,-1.9 -7,-2.7 -2,-0.3 2,-0.3 -0.971 10.1-151.9-144.3 162.0 35.3 2.9 23.5 77 138 A E E +HI 68 104A 76 27,-2.8 26,-2.1 -2,-0.3 27,-1.6 -0.979 28.0 147.9-131.4 150.9 35.9 5.0 20.4 78 139 A G E -HI 67 102A 8 -11,-2.6 -11,-2.3 -2,-0.3 2,-0.4 -0.915 43.6 -84.5-162.8-174.6 33.8 5.2 17.3 79 140 A E E -HI 66 101A 45 22,-1.9 22,-2.9 -2,-0.3 -13,-0.2 -0.919 39.7-127.4-112.2 133.0 32.4 7.0 14.3 80 141 A L E + I 0 100A 6 -15,-3.2 -17,-2.2 -2,-0.4 -16,-0.5 -0.536 44.1 135.9 -81.2 143.0 29.3 9.2 14.6 81 142 A G E -HI 62 99A 2 18,-1.5 18,-3.0 -19,-0.3 2,-0.3 -0.924 41.6-116.3-161.2-176.6 26.3 8.7 12.3 82 143 A T E +HI 61 98A 19 -21,-2.4 -21,-2.6 -2,-0.3 2,-0.3 -0.953 33.1 154.4-131.1 156.8 22.6 8.4 12.2 83 144 A D E - I 0 97A 0 14,-1.8 14,-2.6 -2,-0.3 2,-0.3 -0.965 47.0 -74.7-165.3 172.7 20.2 5.6 11.2 84 145 A L E - I 0 96A 46 -2,-0.3 -63,-2.6 12,-0.2 2,-0.4 -0.606 47.3-171.6 -81.8 138.7 16.7 4.2 11.8 85 146 A V E +DI 20 95A 1 10,-2.8 10,-2.2 -2,-0.3 2,-0.3 -0.973 8.2 178.6-133.5 143.3 16.1 2.5 15.1 86 147 A S E -DI 19 94A 24 -67,-2.0 -67,-2.3 -2,-0.4 8,-0.1 -0.941 24.2-140.9-138.0 164.7 13.3 0.4 16.5 87 148 A I > - 0 0 8 4,-0.6 3,-2.4 6,-0.6 -69,-0.2 -0.888 12.8-161.9-126.6 98.4 12.5 -1.5 19.7 88 149 A P T 3 S+ 0 0 36 0, 0.0 -1,-0.1 0, 0.0 -70,-0.1 0.833 92.7 32.5 -52.7 -40.0 10.6 -4.7 18.8 89 150 A H T 3 S+ 0 0 95 -72,-0.4 -84,-0.1 -84,-0.1 3,-0.1 0.034 118.5 61.1-108.4 25.6 9.2 -5.2 22.3 90 151 A G S < S- 0 0 1 -3,-2.4 75,-0.1 1,-0.3 -87,-0.0 -0.333 109.8 -22.3-118.5-150.5 8.9 -1.5 23.2 91 152 A P - 0 0 21 0, 0.0 2,-1.9 0, 0.0 -4,-0.6 -0.323 67.7-119.4 -56.0 141.1 6.9 1.2 21.6 92 153 A Q S S+ 0 0 127 -3,-0.1 2,-0.3 -6,-0.1 -6,-0.0 -0.498 74.2 109.7 -86.2 70.1 6.0 0.3 18.0 93 154 A V - 0 0 47 -2,-1.9 -6,-0.6 -6,-0.2 2,-0.4 -0.920 57.5-140.1-136.3 162.6 7.7 3.2 16.3 94 155 A T E + I 0 86A 91 -2,-0.3 2,-0.3 -8,-0.1 -8,-0.2 -0.995 24.6 179.0-124.9 127.0 10.8 3.9 14.2 95 156 A V E - 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I 0 76A 25 -2,-0.3 2,-0.4 -29,-0.2 -29,-0.2 -0.987 21.3-148.1-155.3 160.1 37.3 -0.6 22.7 106 167 A D E S- I 0 75A 75 -31,-2.8 -31,-1.6 -2,-0.3 -2,-0.0 -0.935 79.9 -24.8-138.9 113.5 39.0 -1.5 25.9 107 168 A K S S+ 0 0 168 -2,-0.4 -31,-0.1 -33,-0.2 3,-0.0 0.794 106.6 107.8 58.5 29.9 37.5 -3.5 28.8 108 169 A F + 0 0 14 -33,-0.2 -32,-0.1 -34,-0.2 -1,-0.1 0.852 60.5 61.0 -98.3 -63.8 34.0 -2.4 27.6 109 170 A F S S- 0 0 12 7,-0.1 2,-0.5 1,-0.1 -2,-0.1 -0.392 79.4-139.3 -66.8 143.9 32.5 -5.5 26.1 110 171 A I > - 0 0 40 5,-0.2 3,-2.0 -63,-0.1 4,-0.3 -0.912 20.3-107.1-113.8 130.9 32.1 -8.4 28.5 111 172 A Q T 3 S- 0 0 143 -2,-0.5 -64,-0.1 1,-0.3 -63,-0.1 -0.327 102.5 -1.7 -57.3 116.8 32.8 -12.0 27.6 112 173 A G T 3 S+ 0 0 25 -65,-0.3 -1,-0.3 -2,-0.2 -64,-0.1 0.668 90.0 128.4 81.4 18.7 29.4 -13.9 27.3 113 174 A S < - 0 0 1 -3,-2.0 -104,-0.2 -66,-0.2 -2,-0.1 0.728 62.4-137.5 -81.6 -23.4 27.3 -10.9 28.3 114 175 A N + 0 0 50 -4,-0.3 2,-0.3 -67,-0.3 -86,-0.1 0.445 67.0 106.7 76.4 3.6 24.9 -11.2 25.3 115 176 A W - 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