@text "THE DISULFIDE BRIDGE -- STRUCTURAL STABILIZATION OF PROTEINS" Ribonuclease A, a small single-subunit enzyme that cuts RNA, is an excellent example of a protein held together by disulfide bridges (from PDB file, 5RSA). Each of the bridges is the result of two cysteine sidechains in close proximity (as a result of protein folding) undergoing covalent attachment through the terminal sulfurs; the chemical reaction can be depicted as follows: -S-H + H-S- => -S-S-. This reaction occurs under oxidative conditions, typically found in extracellular regions. One or more disulfide bridges can be found to connect domains within a protein (intramolecular bridges). They can also be found in multi-subunit proteins between individual polypeptide chains (intermolecular bridges). The disulfide bridge is critical for maintaining the active conformation in many proteins. In fact, breaking disulfide bonds through reductive processes yields a protein with significantly lower activity. @kinemage 1 @caption Calpha trace of RNase A, with the four disulfide bridges highlighted. Cysteine sulfurs are depicted as gold balls; carbons are green balls. Select view2 for a close-up of one disulfide bond, showing the connection between two sequentially-removed portions of the protein. @onewidth @viewid {4 S-S} @zoom 1.00 @zslab 170 @center 30.267 16.026 13.157 @matrix -0.812147 0.424353 0.400427 0.385290 0.905445 -0.178101 -0.438142 0.009636 -0.898854 @2viewid {Close-up} @2zoom 2.28 @2zslab 200 @2center 20.048 4.269 7.736 @2matrix 0.04866 0.22040 0.97419 -0.05760 0.97435 -0.21756 -0.99715 -0.04553 0.06011 @group {RNase} @subgroup {Calphas} dominant @vectorlist {ca-ca} color=bluetint {ca lys 1}P 17.372 25.599 -1.318 {ca lys 1}17.372 25.599 -1.318 {ca glu 2}19.195 24.738 1.949 {ca thr 3}22.383 26.747 2.254 {ca ala 4}25.655 25.012 3.159 {ca ala 5}25.736 27.038 6.419 {ca ala 6}22.196 25.871 7.429 {ca lys 7}23.094 22.309 6.327 {ca phe 8}26.126 22.443 8.706 {ca glu 9}23.823 23.393 11.561 {ca arg 10}21.305 20.65 10.749 {ca gln 11}23.953 17.905 10.428 {ca his 12}26.354 18.986 13.14 {ca met 13}24.937 21.262 15.889 {ca asp 14}23.324 20.085 19.089 {ca ser 15}23.317 23.068 21.463 {ca ser 16}20.431 21.717 23.489 {ca thr 17}22.3 18.924 25.178 {ca ser 18}25.724 19.105 26.864 {ca ala 19}26.624 15.565 25.829 {ca ala 20}24.865 12.381 24.626 {ca ser 21}23.329 10.656 27.632 {ca ser 22}22.353 7.236 26.351 {ca ser 23}23.626 4.814 23.731 {ca asn 24}20.527 5.723 21.601 {ca tyr 25}21.579 9.308 21. {ca cys 26}23.014 8.67 17.523 {ca asn 27}20.094 6.558 16.399 {ca gln 28}17.684 9.242 17.476 {ca met 29}19.612 12.297 16.155 {ca met 30}20.678 10.819 12.817 {ca lys 31}17.079 10.067 12.038 {ca ser 32}15.551 13.273 13.405 {ca arg 33}18.032 15.515 11.515 {ca asn 34}17.073 13.558 8.4 {ca leu 35}20.597 12.158 7.798 {ca thr 36}19.289 8.698 7.014 {ca lys 37}16.628 9.595 4.496 {ca asp 38}18.199 8.483 1.26 {ca arg 39}20.803 6.147 2.77 {ca cys 40}22.661 5.563 6.06 {ca lys 41}25.326 8.318 6.297 {ca pro 42}28.209 6.182 7.715 {ca val 43}29.703 8.693 10.044 {ca asn 44}28.6 11.954 11.588 {ca thr 45}29.66 14.248 14.475 {ca phe 46}27.37 16.426 16.627 {ca val 47}28.781 19.448 18.564 {ca his 48}27.264 20.375 21.971 {ca glu 49}28.155 24.015 22.153 {ca ser 50}26.545 27.276 21.16 {ca leu 51}26.477 27.996 17.474 {ca ala 52}28.243 31.265 18.166 {ca asp 53}31.217 29.382 19.714 {ca val 54}31.306 27.078 16.71 {ca gln 55}31.102 29.939 14.152 {ca ala 56}33.979 31.61 16.001 {ca val 57}36.263 28.738 15.015 {ca cys 58}36.355 30.441 11.593 {ca ser 59}38.661 32.967 13.126 {ca gln 60}40.865 30.508 15.078 {ca lys 61}43.771 28.489 13.841 {ca asn 62}44.119 28.064 10.104 {ca val 63}44.984 24.473 9.04 {ca ala 64}44.908 22.532 5.726 {ca cys 65}41.706 20.374 5.582 {ca lys 66}42.295 16.648 5.244 {ca asn 67}41.767 16.991 1.441 {ca gly 68}44.475 19.8 1.126 {ca gln 69}42.078 22.81 1.058 {ca thr 70}43.304 25.794 2.987 {ca asn 71}40.023 27.051 4.332 {ca cys 72}39.994 24.79 7.457 {ca tyr 73}40.274 26.059 11.007 {ca gln 74}41.025 24.28 14.3 {ca ser 75}39.372 25.191 17.524 {ca tyr 76}41.739 26.688 20.084 {ca ser 77}39.634 24.841 22.715 {ca thr 78}38.27 21.283 23.228 {ca met 79}34.531 20.902 22.689 {ca ser 80}31.922 18.307 23.794 {ca ile 81}31.23 16.162 20.682 {ca thr 82}29.306 12.947 19.978 {ca asp 83}30.66 10.719 17.189 {ca cys 84}28.149 8.458 15.469 {ca arg 85}29.399 5.472 13.529 {ca glu 86}27.35 2.965 11.598 {ca thr 87}27.128 -0.521 13.258 {ca gly 88}27.443 -3.946 11.608 {ca ser 89}23.734 -4.283 12.167 {ca ser 90}22.876 -1.102 10.179 {ca lys 91}20.42 -1.864 7.36 {ca tyr 92}18.716 0.868 5.308 {ca pro 93}16.13 2.157 6.251 {ca asn 94}17.022 0.873 9.776 {ca cys 95}20.312 2.786 10.06 {ca ala 96}21.986 1.836 13.354 {ca tyr 97}24.785 3.76 15.03 {ca lys 98}27.283 3.427 17.84 {ca thr 99}27.401 6.646 19.974 {ca thr 100}30.772 7.905 21.248 {ca gln 101}30.94 11.023 23.62 {ca ala 102}34.288 12.903 23.746 {ca asn 103}35.891 16.305 24.348 {ca lys 104}38.228 17.262 21.499 {ca his 105}39.259 20.038 19.28 {ca ile 106}37.228 20.423 16.112 {ca ile 107}38.339 21.218 12.514 {ca val 108}35.828 22.861 10.164 {ca ala 109}35.962 24.395 6.635 {ca cys 110}34.607 28.006 6.449 {ca glu 111}33.224 29.981 3.507 {ca gly 112}31.175 33.073 2.538 {ca asn 113}30.36 36.335 4.24 {ca pro 114}29.722 36.156 7.076 {ca tyr 115}32.526 33.511 7.056 {ca val 116}30.914 30.521 8.817 {ca pro 117}31.375 26.739 9.073 {ca val 118}30.091 24.766 6.025 {ca his 119}31.851 21.421 6.622 {ca phe 120}32.927 19.335 9.591 {ca asp 121}36.407 18.112 8.76 {ca ala 122}37.727 16.178 11.842 {ca ser 123}37.937 16.116 15.637 {ca val 124}41.435 16.025 17.085 @subgroup {S-S} dominant @vectorlist {ss} color=yellow {ca cys 65}P 41.706 20.374 5.582 {cb cys 65}40.613 20.618 4.569 {sg cys 65}39.9 22.235 4.768 {sg cys 72}39.222 22.229 6.587 {cb cys 72}40.379 23.319 7.468 {ca cys 72}39.994 24.79 7.457 {ca cys 26}P 23.014 8.67 17.523 {cb cys 26}24.314 7.997 17.133 {sg cys 26}25.715 8.896 17.715 {sg cys 84}26.015 10.222 16.198 {cb cys 84}26.935 9.216 14.986 {ca cys 84}28.149 8.458 15.469 {ca cys 40}P 22.661 5.563 6.06 {cb cys 40}22.867 4.192 6.728 {sg cys 40}21.405 3.143 6.86 {sg cys 95}20.048 4.269 7.736 {cb cys 95}20.502 4.196 9.498 {ca cys 95}20.312 2.786 10.06 {ca cys 58}P 36.355 30.441 11.593 {cb cys 58}35.255 30.831 10.619 {sg cys 58}34.266 29.409 10.161 {sg cys 110}35.415 28.241 9.096 {cb cys 110}35.099 29.042 7.474 {ca cys 110}34.607 28.006 6.449 @balllist {C} radius=.3 color=green {ca cys 65 } P 41.706, 20.374, 5.582 {cb cys 65 } L 40.613, 20.618, 4.569 {cb cys 72 } L 40.379, 23.319, 7.468 {ca cys 72 } L 39.994, 24.790, 7.457 {ca cys 26 } P 23.014, 8.670, 17.523 {cb cys 26 } L 24.314, 7.997, 17.133 {cb cys 84 } L 26.935, 9.216, 14.986 {ca cys 84 } L 28.149, 8.458, 15.469 {ca cys 40 } P 22.661, 5.563, 6.060 {cb cys 40 } L 22.867, 4.192, 6.728 {cb cys 95 } L 20.502, 4.196, 9.498 {ca cys 95 } L 20.312, 2.786, 10.060 {ca cys 58 } P 36.355, 30.441, 11.593 {cb cys 58 } L 35.255, 30.831, 10.619 {cb cys 110 } L 35.099, 29.042, 7.474 {ca cys 110 } L 34.607, 28.006, 6.449 @balllist {S} radius=.3 color=gold { sg cys 65 } L 39.900, 22.235, 4.768 { sg cys 72 } L 39.222, 22.229, 6.587 { sg cys 26 } L 25.715, 8.896, 17.715 { sg cys 84 } L 26.015, 10.222, 16.198 { sg cys 40 } L 21.405, 3.143, 6.860 { sg cys 95 } L 20.048, 4.269, 7.736 { sg cys 58 } L 34.266, 29.409, 10.161 { sg cys 110 } L 35.415, 28.241, 9.096