Curvature-Induced Spatial Ordering of Composition in Lipid Membranes

<div>The effect of spontaneous curvature for different values of <svg height="14.45pt" id="M224" style="vertical-align:-5.48788pt" version="1.1" viewbox="-0.0498162 -8.96212 35.7131 14.45" width="35.7131pt" xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><g transform="matrix(.013,0,0,-0.013,0,0)"><path d="M542 242C542 369 469 429 379 468C370 472 372 496 376 513L416 681L400 691L344 648L219 51C167 69 113 128 113 217C113 324 166 387 266 428L252 459C124 418 23 336 23 202C23 95 90 13 205 -8L182 -132C165 -222 191 -254 209 -261L215 -258L270 -7C415 -4 542 98 542 242ZM469 232C469 126 403 33 279 39L355 409C399 394 469 331 469 232Z" id="g113-243"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="565" vert-adv-y="565"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,7.02,3.132)"><path d="M442 39L431 68C420 63 406 57 394 57C377 57 359 71 359 118V301C359 353 349 393 314 424C291 443 261 451 230 451C171 438 107 401 79 381C59 366 46 355 46 340C46 316 72 295 91 295C106 295 117 303 122 319C131 350 139 371 152 384C163 397 178 404 197 404C246 404 281 365 281 293V278C260 262 186 232 124 213C69 194 41 161 41 112C41 47 92 -12 163 -12C195 -12 243 25 283 51C288 35 293 22 308 7C319 -4 339 -12 354 -12L442 39ZM281 87C262 68 226 51 199 51C169 51 129 74 129 126C129 162 151 181 190 198C211 208 259 229 281 240V87Z" id="g58-95"></path><glyph.data ascent="1007" descent="-369" horiz-adv-x="449" vert-adv-y="449"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,10.932,3.132)"><path d="M487 439H304V409C361 402 367 391 351 344C328 280 297 190 262 110C235 178 192 284 168 353C152 396 153 403 208 409V439H-1V409C54 401 61 394 85 338C133 221 179 111 224 -10H255C312 139 363 260 398 336C426 394 432 401 487 409V439Z" id="g58-116"></path><glyph.data ascent="1007" descent="-369" horiz-adv-x="473" vert-adv-y="473"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,15.291,3.132)"><path d="M384 109C349 76 310 59 268 59C197 59 123 113 120 245C240 248 367 259 384 262C403 266 407 275 407 296C407 376 338 451 254 451H253C201 451 148 425 105 378C65 334 39 272 39 203C39 88 112 -12 235 -12C266 -12 339 5 402 84L384 109ZM230 411C285 411 319 365 319 313C319 298 313 293 294 293C236 291 180 290 125 290C141 370 185 411 230 411Z" id="g58-99"></path><glyph.data ascent="1007" descent="-369" horiz-adv-x="434" vert-adv-y="434"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,19.241,3.132)"><path d="M185 344V453C137 433 92 421 41 413V386C100 378 104 375 104 310V106C104 41 97 35 33 30V0H268V30C190 35 185 41 185 107V283C206 340 240 371 267 371C282 371 296 364 310 351C316 344 325 343 336 348C357 357 369 378 369 399C369 424 345 451 310 451C262 451 219 397 187 344H185Z" id="g58-112"></path><glyph.data ascent="1007" descent="-369" horiz-adv-x="380" vert-adv-y="380"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,22.699,3.132)"><path d="M442 39L431 68C420 63 406 57 394 57C377 57 359 71 359 118V301C359 353 349 393 314 424C291 443 261 451 230 451C171 438 107 401 79 381C59 366 46 355 46 340C46 316 72 295 91 295C106 295 117 303 122 319C131 350 139 371 152 384C163 397 178 404 197 404C246 404 281 365 281 293V278C260 262 186 232 124 213C69 194 41 161 41 112C41 47 92 -12 163 -12C195 -12 243 25 283 51C288 35 293 22 308 7C319 -4 339 -12 354 -12L442 39ZM281 87C262 68 226 51 199 51C169 51 129 74 129 126C129 162 151 181 190 198C211 208 259 229 281 240V87Z" id="g58-95"></path><glyph.data ascent="1007" descent="-369" horiz-adv-x="449" vert-adv-y="449"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,26.739,3.132)"><path d="M472 439C432 432 382 428 334 425C303 442 271 451 236 451H235C158 451 54 398 54 285C54 216 97 171 140 152C124 133 93 105 53 90C52 81 55 63 64 48C77 27 102 5 137 -6C115 -22 77 -53 57 -73C40 -88 32 -107 32 -130C32 -189 96 -253 208 -253C340 -253 461 -159 461 -56C461 44 378 63 314 63C281 63 246 62 207 62C163 62 145 80 145 99C145 114 160 130 175 141C191 138 210 136 226 136C314 136 405 188 405 294C405 329 393 359 376 382L433 378C448 387 469 415 477 430L472 439ZM226 417C282 417 320 364 320 288S281 172 236 171C180 171 142 224 142 300S182 417 226 417ZM248 -9C291 -9 319 -11 346 -22S390 -57 390 -90C390 -151 339 -199 246 -199C171 -199 114 -159 114 -106C114 -79 132 -53 159 -30C178 -15 200 -9 248 -9Z" id="g58-101"></path><glyph.data ascent="1007" descent="-369" horiz-adv-x="479" vert-adv-y="479"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,31.034,3.132)"><path d="M384 109C349 76 310 59 268 59C197 59 123 113 120 245C240 248 367 259 384 262C403 266 407 275 407 296C407 376 338 451 254 451H253C201 451 148 425 105 378C65 334 39 272 39 203C39 88 112 -12 235 -12C266 -12 339 5 402 84L384 109ZM230 411C285 411 319 365 319 313C319 298 313 293 294 293C236 291 180 290 125 290C141 370 185 411 230 411Z" id="g58-99"></path><glyph.data ascent="1007" descent="-369" horiz-adv-x="434" vert-adv-y="434"></glyph.data></g></svg> and sign of the surface curvature. Each plot shows the final (steady state) spatial distribution of the composition field <svg height="12.3916pt" id="M225" style="vertical-align:-3.42948pt" version="1.1" viewbox="-0.0498162 -8.96212 7.47216 12.3916" width="7.47216pt" xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><g transform="matrix(.013,0,0,-0.013,0,0)"><path d="M542 242C542 369 469 429 379 468C370 472 372 496 376 513L416 681L400 691L344 648L219 51C167 69 113 128 113 217C113 324 166 387 266 428L252 459C124 418 23 336 23 202C23 95 90 13 205 -8L182 -132C165 -222 191 -254 209 -261L215 -258L270 -7C415 -4 542 98 542 242ZM469 232C469 126 403 33 279 39L355 409C399 394 469 331 469 232Z" id="g113-243"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="565" vert-adv-y="565"></glyph.data></g></svg>. In each of the four “windows” the left and right plots, (i) and (ii), correspond to <svg height="15.0796pt" id="M226" style="vertical-align:-3.1815pt" version="1.1" viewbox="-0.0498162 -11.8981 45.9481 15.0796" width="45.9481pt" xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><rect height="0.65243" width="10.9609" x="0" y="-11.1958"></rect><g transform="matrix(.013,0,0,-0.013,0,0)"><path d="M828 650H570L564 622C646 614 650 609 634 524L604 366H253L281 524C296 608 308 615 382 622L388 650H132L126 622C211 615 214 609 198 524L121 122C106 42 102 35 23 28L17 0H276L282 28C196 35 191 40 206 122L243 324H597L559 123C544 42 537 35 452 28L446 0H710L716 28C632 35 628 43 643 122L719 524C735 610 741 615 822 622L828 650Z" id="g113-73"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="840" vert-adv-y="840"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,10.961,3.132)"><path d="M418 650H141L133 618C225 611 232 604 218 531L141 120C126 45 119 37 33 32L24 0H301L308 32C220 37 214 44 227 120L305 531C319 605 325 611 411 618L418 650Z" id="g50-74"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="442" vert-adv-y="442"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,19.055,0)"><path d="M535 323V373H52V323H535ZM535 138V188H52V138H535Z" id="g117-34"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="587" vert-adv-y="587"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,30.318,0)"><path d="M241 635C89 635 35 457 35 312C35 153 89 -12 240 -12C390 -12 443 166 443 312C443 466 390 635 241 635ZM238 602C329 602 354 454 354 312C354 172 330 22 240 22C152 22 124 173 124 313S148 602 238 602Z" id="g113-49"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="480" vert-adv-y="480"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,36.558,0)"><path d="M113 -12C146 -12 170 11 170 46C170 78 146 103 114 103S58 78 58 46C58 11 82 -12 113 -12Z" id="g113-47"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="228" vert-adv-y="228"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,39.522,0)"><path d="M384 0V27C293 34 287 42 287 114V635C232 613 172 594 109 583V559L157 557C201 555 205 550 205 499V114C205 42 199 34 109 27V0H384Z" id="g113-50"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="480" vert-adv-y="480"></glyph.data></g></svg> and <svg height="15.0796pt" id="M227" style="vertical-align:-3.1815pt" version="1.1" viewbox="-0.0498162 -11.8981 36.7096 15.0796" width="36.7096pt" xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><rect height="0.65243" width="10.9609" x="0" y="-11.1958"></rect><g transform="matrix(.013,0,0,-0.013,0,0)"><path d="M828 650H570L564 622C646 614 650 609 634 524L604 366H253L281 524C296 608 308 615 382 622L388 650H132L126 622C211 615 214 609 198 524L121 122C106 42 102 35 23 28L17 0H276L282 28C196 35 191 40 206 122L243 324H597L559 123C544 42 537 35 452 28L446 0H710L716 28C632 35 628 43 643 122L719 524C735 610 741 615 822 622L828 650Z" id="g113-73"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="840" vert-adv-y="840"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,10.961,3.132)"><path d="M418 650H141L133 618C225 611 232 604 218 531L141 120C126 45 119 37 33 32L24 0H301L308 32C220 37 214 44 227 120L305 531C319 605 325 611 411 618L418 650Z" id="g50-74"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="442" vert-adv-y="442"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,19.055,0)"><path d="M535 323V373H52V323H535ZM535 138V188H52V138H535Z" id="g117-34"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="587" vert-adv-y="587"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,30.318,0)"><path d="M384 0V27C293 34 287 42 287 114V635C232 613 172 594 109 583V559L157 557C201 555 205 550 205 499V114C205 42 199 34 109 27V0H384Z" id="g113-50"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="480" vert-adv-y="480"></glyph.data></g></svg>, respectively. <svg height="8.55521pt" id="M228" style="vertical-align:-0.2063904pt" version="1.1" viewbox="-0.0498162 -8.34882 31.5818 8.55521" width="31.5818pt" xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><g transform="matrix(.013,0,0,-0.013,0,0)"><path d="M766 88L752 113C719 83 690 64 681 64C674 64 672 74 679 103L724 292C758 436 724 448 701 448C680 448 666 442 639 429C594 407 514 350 441 252H439L447 289C476 423 450 448 419 448C398 448 379 441 355 427C307 400 234 344 162 249H160L180 324C203 409 197 448 170 448C144 448 82 413 23 349L35 321C57 343 96 374 108 374C115 374 117 371 111 341C87 227 57 112 24 -6L32 -12C53 -4 81 4 108 6C119 68 134 128 149 171C177 229 309 383 364 383C387 383 388 355 373 282C354 190 330 92 303 -6L309 -12C332 -4 356 3 386 6C396 63 411 122 424 171C458 236 590 383 642 383C658 383 664 369 652 315L603 91C587 20 593 -12 619 -12C642 -12 708 23 766 88Z" id="g113-110"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="789" vert-adv-y="789"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,13.889,0)"><path d="M535 323V373H52V323H535ZM535 138V188H52V138H535Z" id="g117-34"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="587" vert-adv-y="587"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,25.152,0)"><path d="M412 140C382 77 369 73 315 73H129L270 222C362 320 402 379 402 466C402 571 322 635 234 635C177 635 130 609 99 576L42 495L64 475C90 514 133 568 201 568C274 568 318 519 318 435C318 349 255 267 193 193C144 135 87 78 32 23V0H405C417 45 427 89 440 131L412 140Z" id="g113-51"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="480" vert-adv-y="480"></glyph.data></g></svg>, <svg height="12.4698pt" id="M229" style="vertical-align:-3.18147pt" version="1.1" viewbox="-0.0498162 -9.28833 40.4882 12.4698" width="40.4882pt" xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><g transform="matrix(.013,0,0,-0.013,0,0)"><path d="M480 416C480 431 465 448 438 448C388 448 312 383 252 330C217 299 188 273 155 237H153L257 680C262 700 263 712 253 712C240 712 183 684 97 674L92 648L126 647C166 646 172 645 163 606L23 -6L29 -12C51 -5 77 2 107 8C115 62 130 128 142 180C153 193 179 220 204 241C231 170 259 106 288 54C317 0 336 -12 358 -12C381 -12 423 2 477 80L460 100C434 74 408 54 398 54C385 54 374 65 351 107C326 154 282 241 263 299C296 332 351 377 403 377C424 377 436 372 445 368C449 366 456 368 462 375C472 386 480 402 480 416Z" id="g113-108"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="503" vert-adv-y="503"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,6.474,3.132)"><path d="M418 650H141L133 618C225 611 232 604 218 531L141 120C126 45 119 37 33 32L24 0H301L308 32C220 37 214 44 227 120L305 531C319 605 325 611 411 618L418 650Z" id="g50-74"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="442" vert-adv-y="442"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,14.568,0)"><path d="M535 323V373H52V323H535ZM535 138V188H52V138H535Z" id="g117-34"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="587" vert-adv-y="587"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,25.831,0)"><path d="M480 416C480 431 465 448 438 448C388 448 312 383 252 330C217 299 188 273 155 237H153L257 680C262 700 263 712 253 712C240 712 183 684 97 674L92 648L126 647C166 646 172 645 163 606L23 -6L29 -12C51 -5 77 2 107 8C115 62 130 128 142 180C153 193 179 220 204 241C231 170 259 106 288 54C317 0 336 -12 358 -12C381 -12 423 2 477 80L460 100C434 74 408 54 398 54C385 54 374 65 351 107C326 154 282 241 263 299C296 332 351 377 403 377C424 377 436 372 445 368C449 366 456 368 462 375C472 386 480 402 480 416Z" id="g113-108"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="503" vert-adv-y="503"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,32.305,3.132)"><path d="M418 650H141L133 618C225 611 232 604 218 531L141 120C126 45 119 37 33 32L24 0H301L308 32C220 37 214 44 227 120L305 531C319 605 325 611 411 618L418 650Z" id="g50-74"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="442" vert-adv-y="442"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,35.836,3.132)"><path d="M418 650H141L133 618C225 611 232 604 218 531L141 120C126 45 119 37 33 32L24 0H301L308 32C220 37 214 44 227 120L305 531C319 605 325 611 411 618L418 650Z" id="g50-74"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="442" vert-adv-y="442"></glyph.data></g></svg>, and <svg height="11.8174pt" id="M230" style="vertical-align:-3.1815pt" version="1.1" viewbox="-0.0498162 -8.6359 55.3768 11.8174" width="55.3768pt" xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><g transform="matrix(.013,0,0,-0.013,0,0)"><path d="M828 650H570L564 622C646 614 650 609 634 524L604 366H253L281 524C296 608 308 615 382 622L388 650H132L126 622C211 615 214 609 198 524L121 122C106 42 102 35 23 28L17 0H276L282 28C196 35 191 40 206 122L243 324H597L559 123C544 42 537 35 452 28L446 0H710L716 28C632 35 628 43 643 122L719 524C735 610 741 615 822 622L828 650Z" id="g113-73"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="840" vert-adv-y="840"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,10.075,3.132)"><path d="M418 650H141L133 618C225 611 232 604 218 531L141 120C126 45 119 37 33 32L24 0H301L308 32C220 37 214 44 227 120L305 531C319 605 325 611 411 618L418 650Z" id="g50-74"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="442" vert-adv-y="442"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,13.606,3.132)"><path d="M418 650H141L133 618C225 611 232 604 218 531L141 120C126 45 119 37 33 32L24 0H301L308 32C220 37 214 44 227 120L305 531C319 605 325 611 411 618L418 650Z" id="g50-74"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="442" vert-adv-y="442"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,21.713,0)"><path d="M535 323V373H52V323H535ZM535 138V188H52V138H535Z" id="g117-34"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="587" vert-adv-y="587"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,32.976,0)"><path d="M535 230V280H52V230H535Z" id="g117-33"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="587" vert-adv-y="587"></glyph.data></g><g transform="matrix(.013,0,0,-0.013,40.636,0)"><path d="M828 650H570L564 622C646 614 650 609 634 524L604 366H253L281 524C296 608 308 615 382 622L388 650H132L126 622C211 615 214 609 198 524L121 122C106 42 102 35 23 28L17 0H276L282 28C196 35 191 40 206 122L243 324H597L559 123C544 42 537 35 452 28L446 0H710L716 28C632 35 628 43 643 122L719 524C735 610 741 615 822 622L828 650Z" id="g113-73"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="840" vert-adv-y="840"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,50.711,3.132)"><path d="M418 650H141L133 618C225 611 232 604 218 531L141 120C126 45 119 37 33 32L24 0H301L308 32C220 37 214 44 227 120L305 531C319 605 325 611 411 618L418 650Z" id="g50-74"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="442" vert-adv-y="442"></glyph.data></g></svg>.</div>

Computational and Mathematical Methods in Medicine

Curvature-Induced Spatial Ordering of Composition in Lipid Membranes

Figure 6