All Three Rows of Outer Hair Cells Are Required for Cochlear Amplification

<div>Electrical circuit model of the cell membrane. <svg height="11.927pt" id="M220" style="vertical-align:-3.291101pt" version="1.1" viewbox="-0.0498162 -8.6359 16.067 11.927" width="16.067pt" 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="M610 18C585 26 567 34 540 68C517 97 499 128 476 171C452 215 425 276 413 304C496 332 570 394 570 494C570 555 545 595 509 619S419 650 364 650H139L133 622C216 615 219 612 203 527L129 132C112 40 105 36 23 28L17 0H279L285 28C199 34 194 40 211 132L239 284H284C320 284 334 275 351 236C374 182 394 140 420 93C459 23 495 -1 592 -8H600L610 18ZM480 485C480 424 449 372 403 342C374 323 338 316 293 316H245L291 562C296 589 301 601 311 608S337 618 358 618C432 618 480 575 480 485Z" id="g113-83"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="627" vert-adv-y="627"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,8.086,3.132)"><path d="M781 91L765 118C733 86 702 70 693 70C686 70 685 77 692 106L738 297C772 439 736 451 712 451S675 445 647 431C605 410 526 356 452 257H450L459 294C491 426 459 451 429 451C403 451 386 445 361 431C312 404 239 343 170 253H168L188 333C208 411 205 451 175 451C148 451 85 417 24 346L39 318C57 338 100 374 112 374C120 374 122 372 115 339C91 226 62 112 28 -6L37 -12C59 -4 87 3 114 6C125 68 142 127 154 168C184 227 316 383 372 383C397 383 396 366 381 289C362 192 338 92 312 -6L317 -12C339 -5 367 3 396 6L434 170C468 234 598 383 650 383C668 383 676 369 664 319L609 90C593 23 600 -12 629 -12C654 -12 722 23 781 91Z" id="g50-110"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="805" vert-adv-y="805"></glyph.data></g></svg> and <svg height="11.927pt" id="M221" style="vertical-align:-3.291101pt" version="1.1" viewbox="-0.0498162 -8.6359 16.602 11.927" width="16.602pt" 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="M645 631C614 643 545 666 457 666C215 666 23 519 23 283C23 90 158 -16 337 -16C412 -16 489 2 522 10C543 39 590 127 606 167L580 181C519 89 459 18 348 18C201 18 122 136 122 287C122 464 244 632 435 632C544 632 602 595 608 472L639 475C643 526 645 581 645 631Z" id="g113-68"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="668" vert-adv-y="668"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,8.619,3.132)"><path d="M781 91L765 118C733 86 702 70 693 70C686 70 685 77 692 106L738 297C772 439 736 451 712 451S675 445 647 431C605 410 526 356 452 257H450L459 294C491 426 459 451 429 451C403 451 386 445 361 431C312 404 239 343 170 253H168L188 333C208 411 205 451 175 451C148 451 85 417 24 346L39 318C57 338 100 374 112 374C120 374 122 372 115 339C91 226 62 112 28 -6L37 -12C59 -4 87 3 114 6C125 68 142 127 154 168C184 227 316 383 372 383C397 383 396 366 381 289C362 192 338 92 312 -6L317 -12C339 -5 367 3 396 6L434 170C468 234 598 383 650 383C668 383 676 369 664 319L609 90C593 23 600 -12 629 -12C654 -12 722 23 781 91Z" id="g50-110"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="805" vert-adv-y="805"></glyph.data></g></svg> are the membrane resistance and membrane capacitance, respectively. <svg height="11.927pt" id="M222" style="vertical-align:-3.291101pt" version="1.1" viewbox="-0.0498162 -8.6359 12.6874 11.927" width="12.6874pt" 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="M405 650H141L135 622C222 616 230 610 215 535L133 116C118 41 113 33 29 28L23 0H289L295 28C209 33 205 40 219 116L298 535C312 609 317 616 399 622L405 650Z" id="g113-74"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="428" vert-adv-y="428"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,4.719,3.132)"><path d="M781 91L765 118C733 86 702 70 693 70C686 70 685 77 692 106L738 297C772 439 736 451 712 451S675 445 647 431C605 410 526 356 452 257H450L459 294C491 426 459 451 429 451C403 451 386 445 361 431C312 404 239 343 170 253H168L188 333C208 411 205 451 175 451C148 451 85 417 24 346L39 318C57 338 100 374 112 374C120 374 122 372 115 339C91 226 62 112 28 -6L37 -12C59 -4 87 3 114 6C125 68 142 127 154 168C184 227 316 383 372 383C397 383 396 366 381 289C362 192 338 92 312 -6L317 -12C339 -5 367 3 396 6L434 170C468 234 598 383 650 383C668 383 676 369 664 319L609 90C593 23 600 -12 629 -12C654 -12 722 23 781 91Z" id="g50-110"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="805" vert-adv-y="805"></glyph.data></g></svg> is the current across the membrane and <svg height="11.927pt" id="M223" style="vertical-align:-3.291101pt" version="1.1" viewbox="-0.0498162 -8.6359 15.3102 11.927" width="15.3102pt" 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="M697 650H468L461 623L492 619C539 613 547 605 518 546C481 471 367 264 278 116H276C239 278 197 500 186 567C180 604 185 613 226 619L252 623L260 650H24L17 623C78 617 92 613 108 533L216 -11H247C365 200 515 462 560 529C616 612 624 615 689 623L697 650Z" id="g113-87"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="709" vert-adv-y="709"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,7.332,3.132)"><path d="M781 91L765 118C733 86 702 70 693 70C686 70 685 77 692 106L738 297C772 439 736 451 712 451S675 445 647 431C605 410 526 356 452 257H450L459 294C491 426 459 451 429 451C403 451 386 445 361 431C312 404 239 343 170 253H168L188 333C208 411 205 451 175 451C148 451 85 417 24 346L39 318C57 338 100 374 112 374C120 374 122 372 115 339C91 226 62 112 28 -6L37 -12C59 -4 87 3 114 6C125 68 142 127 154 168C184 227 316 383 372 383C397 383 396 366 381 289C362 192 338 92 312 -6L317 -12C339 -5 367 3 396 6L434 170C468 234 598 383 650 383C668 383 676 369 664 319L609 90C593 23 600 -12 629 -12C654 -12 722 23 781 91Z" id="g50-110"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="805" vert-adv-y="805"></glyph.data></g></svg> is the membrane potential. The membrane impedance, which acts as the low-pass filter, consists of a parallel connection of <svg height="11.927pt" id="M224" style="vertical-align:-3.291101pt" version="1.1" viewbox="-0.0498162 -8.6359 16.067 11.927" width="16.067pt" 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="M610 18C585 26 567 34 540 68C517 97 499 128 476 171C452 215 425 276 413 304C496 332 570 394 570 494C570 555 545 595 509 619S419 650 364 650H139L133 622C216 615 219 612 203 527L129 132C112 40 105 36 23 28L17 0H279L285 28C199 34 194 40 211 132L239 284H284C320 284 334 275 351 236C374 182 394 140 420 93C459 23 495 -1 592 -8H600L610 18ZM480 485C480 424 449 372 403 342C374 323 338 316 293 316H245L291 562C296 589 301 601 311 608S337 618 358 618C432 618 480 575 480 485Z" id="g113-83"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="627" vert-adv-y="627"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,8.086,3.132)"><path d="M781 91L765 118C733 86 702 70 693 70C686 70 685 77 692 106L738 297C772 439 736 451 712 451S675 445 647 431C605 410 526 356 452 257H450L459 294C491 426 459 451 429 451C403 451 386 445 361 431C312 404 239 343 170 253H168L188 333C208 411 205 451 175 451C148 451 85 417 24 346L39 318C57 338 100 374 112 374C120 374 122 372 115 339C91 226 62 112 28 -6L37 -12C59 -4 87 3 114 6C125 68 142 127 154 168C184 227 316 383 372 383C397 383 396 366 381 289C362 192 338 92 312 -6L317 -12C339 -5 367 3 396 6L434 170C468 234 598 383 650 383C668 383 676 369 664 319L609 90C593 23 600 -12 629 -12C654 -12 722 23 781 91Z" id="g50-110"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="805" vert-adv-y="805"></glyph.data></g></svg> and <svg height="11.927pt" id="M225" style="vertical-align:-3.291101pt" version="1.1" viewbox="-0.0498162 -8.6359 16.602 11.927" width="16.602pt" 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="M645 631C614 643 545 666 457 666C215 666 23 519 23 283C23 90 158 -16 337 -16C412 -16 489 2 522 10C543 39 590 127 606 167L580 181C519 89 459 18 348 18C201 18 122 136 122 287C122 464 244 632 435 632C544 632 602 595 608 472L639 475C643 526 645 581 645 631Z" id="g113-68"></path><glyph.data ascent="3473" descent="-2876" horiz-adv-x="668" vert-adv-y="668"></glyph.data></g><g transform="matrix(.0091,0,0,-0.0091,8.619,3.132)"><path d="M781 91L765 118C733 86 702 70 693 70C686 70 685 77 692 106L738 297C772 439 736 451 712 451S675 445 647 431C605 410 526 356 452 257H450L459 294C491 426 459 451 429 451C403 451 386 445 361 431C312 404 239 343 170 253H168L188 333C208 411 205 451 175 451C148 451 85 417 24 346L39 318C57 338 100 374 112 374C120 374 122 372 115 339C91 226 62 112 28 -6L37 -12C59 -4 87 3 114 6C125 68 142 127 154 168C184 227 316 383 372 383C397 383 396 366 381 289C362 192 338 92 312 -6L317 -12C339 -5 367 3 396 6L434 170C468 234 598 383 650 383C668 383 676 369 664 319L609 90C593 23 600 -12 629 -12C654 -12 722 23 781 91Z" id="g50-110"></path><glyph.data ascent="3443" descent="-2856" horiz-adv-x="805" vert-adv-y="805"></glyph.data></g></svg>.</div>

BioMed Research International

fig15

Figure 15

Figure 15: All Three Rows of Outer Hair Cells Are Required for Cochlear Amplification 