3.06:_Golgi_Apparatus

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<p class="lt-bio-3970" style="background-color: unset;">The Golgi apparatus is a cell structure mainly devoted to processing the proteins synthesized in the endoplasmic reticulum (ER). Some of these will eventually end up as integral membrane proteins embedded in the plasma membrane. Other proteins moving through the Golgi will end up in lysosomes or be secreted by exocytosis (e.g., digestive enzymes). The major processing activity is <strong>glycosylation</strong>: the adding of sugar molecules to form <strong>glycoproteins</strong>. In some cells, e.g., mucus-secreting cells in epithelia, the amount of carbohydrate so far exceeds that of the protein that the product is called a <strong>mucopolysaccharide</strong> (also known as a <strong>proteoglycan</strong>). In plant cells, the Golgi secretes the cell plate and cell wall.</p>

<p class="lt-bio-3970" style="background-color: unset;">Small peptides, e.g., some <strong>hormones</strong> and <strong>neurotransmitters</strong>, are typically too small to be synthesized directly by ribosomes. Instead, the ribosomes on the ER synthesize a large precursor protein that is later cut up into small peptide fragments as it traverses the Golgi. For example, proopiomelanocortin (POMC) is a polypeptide of 241 amino acids from which is cut ACTH, alpha and beta MSH, beta- endorphin, and others.  POMC is cleaved to give rise to multiple peptide hormones.</p>

<ul>
    <li class="lt-bio-3970" style="background-color: unset;">&alpha;-MSH produced by neurons in the arcuate nucleus has important roles in the regulation of appetite and sexual behavior, while &alpha;-MSH secreted from the intermediate lobe of the pituitary regulates the production of melanin.</li>
    <li class="lt-bio-3970" style="background-color: unset;">ACTH is a peptide hormone that regulates the secretion of glucocorticoids from the adrenal cortex.</li>
    <li class="lt-bio-3970" style="background-color: unset;">&beta;-Endorphin and [Met]enkephalin are endogenous opioid peptides with widespread actions in the brain.</li>
</ul>

<figure><img style="width: 602px; height: 336px;" class="internal" alt="Flowchart of POMC cleavage into ACTH, MSH peptides, lipotropins, CLIP, and beta-endorphin." loading="lazy" width="1280" height="717" src="https://bio.libretexts.org/@api/deki/files/5588/POMC.png?revision=1&amp;size=bestfit&amp;width=602&amp;height=336" aria-describedby="remedy-description-f76310786564bcad" /><span id="remedy-description-f76310786564bcad" data-remedy-image-description="f76310786564bcad" style="display: block; white-space: pre-line;">A hierarchical diagram showing how the precursor protein POMC is cut into smaller peptide hormones. At the top, a wide box labeled 'POMC' spans the full width. Dashed vertical lines drop from POMC to three second-level boxes, left to right: '??-MSH' (shaded gray), 'ACTH' (shaded gray), and '??-lipotropin' (unshaded); the dashed lines align with the segment of POMC that yields each product. ??-MSH is a terminal product. From ACTH, dashed lines lead down to two third-level terminal boxes: '??-MSH' (shaded gray) and 'CLIP' (unshaded). From ??-lipotropin, dashed lines lead to two third-level terminal boxes: '??-lipotropin' (unshaded) and '??-endorphin' (unshaded). From ??-lipotropin, dashed lines lead down to a fourth-level terminal box: '??-MSH' (shaded gray). The gray shading distinguishes the melanocortin peptides (??-MSH, ACTH, ??-MSH, ??-MSH) from the other cleavage products (??-lipotropin, CLIP, ??-lipotropin, ??-endorphin), which are unshaded. Each dashed connector indicates that the lower peptide is cleaved from the corresponding portion of the peptide above it.</span><figcaption>Figure <mjx-container class="MathJax CtxtMenu_Attached_0" jax="SVG" overflow="linebreak" tabindex="0" ctxtmenu_counter="96" style="font-size: 85%; position: relative;"><svg width="4.652ex" height="1.557ex" role="img" focusable="false" viewbox="0 -666 2056 688" aria-hidden="true" style="vertical-align: -0.05ex;"><defs><path id="MJX-97-NCM-N-33" d="M303 353C369 378 431 441 431 526C431 569 410 604 369 631C333 654 292 666 246 666C201 666 162 654 127 631C88 605 68 571 68 528C68 495 90 472 122 472C154 472 176 495 176 527C176 560 157 578 119 580C145 615 186 633 242 633C302 633 332 598 332 527C332 485 324 450 309 421C282 373 245 364 183 364C171 362 165 357 165 348C165 333 172 333 192 333L235 333C310 333 348 280 348 173C348 88 317 14 241 14C176 14 128 36 99 80C134 79 160 105 160 139C160 173 135 198 101 198C62 198 42 178 42 137C42 88 64 49 108 18C147-9 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display="inline"><math data-latex="\PageIndex{1}" data-semantic-structure="(5 0 1 2 3 4)"><mrow data-mjx-texclass="ORD" data-latex="1}" data-semantic-type="punctuated" data-semantic-role="sequence" data-semantic-annotation="depth:1" data-semantic-="" data-semantic-children="0,1,2,3,4" data-semantic-content="1,3" data-semantic-attributes="latex:\PageIndex{1};texclass:ORD" data-semantic-owns="0 1 2 3 4" data-semantic-level-number="0" data-speech-node="true"><mn data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:3" data-semantic-level-number="1" data-speech-node="true">3</mn><mrow data-mjx-texclass="ORD" data-latex="{.}"><mo data-latex="." data-semantic-type="punctuation" data-semantic-role="fullstop" data-semantic-annotation="nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:{.};texclass:ORD" data-semantic-operator="punctuated" data-semantic-level-number="1" data-speech-node="true">.</mo></mrow><mn data-latex="6" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:6" data-semantic-level-number="1" data-speech-node="true">6</mn><mrow data-mjx-texclass="ORD" data-latex="{.}"><mo data-latex="." data-semantic-type="punctuation" data-semantic-role="fullstop" data-semantic-annotation="nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:{.};texclass:ORD" data-semantic-operator="punctuated" data-semantic-level-number="1" data-speech-node="true">.</mo></mrow><mn data-latex="1" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:1" data-semantic-level-number="1" data-speech-node="true">1</mn></mrow></math></mjx-assistive-mml></mjx-container>: Proopiomelanocortin (POMC) and Melanocortin Peptides.</figcaption></figure>

<p class="lt-bio-3970" style="background-color: unset;">The Golgi consists of a stack of membrane-bounded <strong>cisternae</strong> located between the endoplasmic reticulum and the cell surface (Figure <mjx-container class="MathJax CtxtMenu_Attached_0" jax="SVG" overflow="linebreak" tabindex="0" ctxtmenu_counter="97" style="font-size: 85%; position: relative;"><svg width="4.652ex" height="1.557ex" role="img" focusable="false" viewbox="0 -666 2056 688" aria-hidden="true" style="vertical-align: -0.05ex;"><defs><path id="MJX-98-NCM-N-33" d="M303 353C369 378 431 441 431 526C431 569 410 604 369 631C333 654 292 666 246 666C201 666 162 654 127 631C88 605 68 571 68 528C68 495 90 472 122 472C154 472 176 495 176 527C176 560 157 578 119 580C145 615 186 633 242 633C302 633 332 598 332 527C332 485 324 450 309 421C282 373 245 364 183 364C171 362 165 357 165 348C165 333 172 333 192 333L235 333C310 333 348 280 348 173C348 88 317 14 241 14C176 14 128 36 99 80C134 79 160 105 160 139C160 173 135 198 101 198C62 198 42 178 42 137C42 88 64 49 108 18C147-9 193-22 244-22C301-22 350-3 393 34C436 71 457 117 457 173C457 267 383 332 303 353Z"></path><path id="MJX-98-NCM-N-2E" d="M192 53C192 82 168 106 139 106C110 106 86 82 86 53C86 24 110 0 139 0C168 0 192 24 192 53Z"></path><path id="MJX-98-NCM-N-36" d="M383 504C416 504 432 521 432 555C432 627 378 666 304 666C221 666 155 627 106 548C63 480 42 403 42 316C42 189 65 100 112 47C152 1 198-22 251-22C312-22 362 1 401 47C438 91 457 144 457 205C457 266 439 318 403 361C365 407 316 431 257 431C205 431 165 402 138 346L138 352C138 465 166 561 226 605C252 624 279 633 306 633C342 633 368 623 385 602C351 602 334 583 334 553C334 525 355 504 383 504M344 340C355 317 361 272 361 206C361 141 356 98 345 76C325 35 294 14 251 14C222 14 200 24 184 44C171 60 162 74 158 85C146 116 140 163 140 227C140 255 144 282 151 308C164 355 201 399 256 399C295 399 325 379 344 340Z"></path><path id="MJX-98-NCM-N-32" d="M237 666C186 666 143 648 106 612C69 576 50 534 50 483C50 449 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data-speech-node="true">.</mo></mrow><mn data-latex="2" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:2" data-semantic-level-number="1" data-speech-node="true">2</mn></mrow></math></mjx-assistive-mml></mjx-container>). Many different enzymes (proteins) are present in the Golgi to perform its various synthetic activities. So there must be mechanisms to sort out the processed proteins and send them on to their destinations while reclaiming processing proteins (e.g., glycosylases) for reuse.</p>

<figure><img class="internal" alt="Electron micrograph with two Golgi stacks outlined in red." loading="lazy" src="https://bio.libretexts.org/@api/deki/files/5583/Golgi.jpg?revision=1" aria-describedby="remedy-description-aee7e05e86935fd8" /><span id="remedy-description-aee7e05e86935fd8" data-remedy-image-description="aee7e05e86935fd8" style="display: block; white-space: pre-line;">Two red rectangles, one near the top and one near the bottom, enclose stacks of flattened, curved membrane compartments. Small rounded vesicles surround their edges. Additional membranes and organelles fill the grainy cytoplasm. There is no scale bar or other label in the image.</span><figcaption>Figure <mjx-container class="MathJax CtxtMenu_Attached_0" jax="SVG" overflow="linebreak" tabindex="0" ctxtmenu_counter="98" style="font-size: 85%; position: relative;"><svg width="4.652ex" height="1.557ex" role="img" focusable="false" viewbox="0 -666 2056 688" aria-hidden="true" style="vertical-align: -0.05ex;"><defs><path id="MJX-99-NCM-N-33" d="M303 353C369 378 431 441 431 526C431 569 410 604 369 631C333 654 292 666 246 666C201 666 162 654 127 631C88 605 68 571 68 528C68 495 90 472 122 472C154 472 176 495 176 527C176 560 157 578 119 580C145 615 186 633 242 633C302 633 332 598 332 527C332 485 324 450 309 421C282 373 245 364 183 364C171 362 165 357 165 348C165 333 172 333 192 333L235 333C310 333 348 280 348 173C348 88 317 14 241 14C176 14 128 36 99 80C134 79 160 105 160 139C160 173 135 198 101 198C62 198 42 178 42 137C42 88 64 49 108 18C147-9 193-22 244-22C301-22 350-3 393 34C436 71 457 117 457 173C457 267 383 332 303 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470C352 403 318 331 250 255L62 43C49 28 50 29 50 0L421 0L450 180L417 180C409 129 402 100 396 91C391 86 361 84 306 84L139 84L236 179C304 243 390 312 419 365C439 400 449 435 449 470C449 588 357 666 237 666Z"></path></defs><g stroke="currentColor" fill="currentColor" stroke-width="0" transform="scale(1,-1)"><g data-mml-node="math" data-latex="\PageIndex{2}" data-semantic-structure="(5 0 1 2 3 4)"><g data-mml-node="TeXAtom" data-mjx-texclass="ORD" data-latex="2}" data-semantic-type="punctuated" data-semantic-role="sequence" data-semantic-annotation="depth:1" data-semantic-id="5" data-semantic-children="0,1,2,3,4" data-semantic-content="1,3" data-semantic-attributes="latex:\PageIndex{2};texclass:ORD" data-semantic-owns="0 1 2 3 4" data-semantic-level-number="0" data-speech-node="true"><g data-mml-node="mn" data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-id="0" 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transform="translate(778,0)"><use data-c="36" href="#MJX-99-NCM-N-36"></use></g><g data-mml-node="TeXAtom" data-mjx-texclass="ORD" data-latex="{.}" transform="translate(1278,0)"><g data-mml-node="mo" data-latex="." data-semantic-type="punctuation" data-semantic-role="fullstop" data-semantic-annotation="nemeth:number;depth:2" data-semantic-id="3" data-semantic-parent="5" data-semantic-attributes="latex:{.};texclass:ORD" data-semantic-operator="punctuated" data-semantic-level-number="1" data-speech-node="true"><use data-c="2E" href="#MJX-99-NCM-N-2E"></use></g></g><g data-mml-node="mn" data-latex="2" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-id="4" data-semantic-parent="5" data-semantic-attributes="latex:2" data-semantic-level-number="1" data-speech-node="true" transform="translate(1556,0)"><use data-c="32" href="#MJX-99-NCM-N-32"></use></g></g></g></g></svg><mjx-assistive-mml unselectable="on" display="inline"><math data-latex="\PageIndex{2}" data-semantic-structure="(5 0 1 2 3 4)"><mrow data-mjx-texclass="ORD" data-latex="2}" data-semantic-type="punctuated" data-semantic-role="sequence" data-semantic-annotation="depth:1" data-semantic-="" data-semantic-children="0,1,2,3,4" data-semantic-content="1,3" data-semantic-attributes="latex:\PageIndex{2};texclass:ORD" data-semantic-owns="0 1 2 3 4" data-semantic-level-number="0" data-speech-node="true"><mn data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:3" data-semantic-level-number="1" data-speech-node="true">3</mn><mrow data-mjx-texclass="ORD" data-latex="{.}"><mo data-latex="." data-semantic-type="punctuation" data-semantic-role="fullstop" data-semantic-annotation="nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:{.};texclass:ORD" data-semantic-operator="punctuated" data-semantic-level-number="1" data-speech-node="true">.</mo></mrow><mn data-latex="6" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:6" data-semantic-level-number="1" data-speech-node="true">6</mn><mrow data-mjx-texclass="ORD" data-latex="{.}"><mo data-latex="." data-semantic-type="punctuation" data-semantic-role="fullstop" data-semantic-annotation="nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:{.};texclass:ORD" data-semantic-operator="punctuated" data-semantic-level-number="1" data-speech-node="true">.</mo></mrow><mn data-latex="2" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:2" data-semantic-level-number="1" data-speech-node="true">2</mn></mrow></math></mjx-assistive-mml></mjx-container>: Golgi apparatus in a bat  cell (Courtesy of Keith R. Porter)</figcaption></figure>

<span id="The_Outbound_Path_(Membrane_Fission)"></span><span id="The_Outbound_Path_(Membrane_Fission)-2"></span><h2 style="background-color: unset;" class="lt-bio-3970">The Outbound Path (Membrane Fission)</h2>

<p class="lt-bio-3970" style="background-color: unset;">Two mechanisms appear to participate in the migration of proteins from the endoplasmic reticulum through the Golgi apparatus.</p>

<ul>
    <li class="lt-bio-3970" style="background-color: unset;"><strong>Mechanism 1:</strong> Transition vesicles pinch off from the surface of the endoplasmic reticulum carrying integral membrane proteins, soluble proteins awaiting processing, and processing enzymes. Pinching off requires that the vesicle be coated with COPII (Coat Protein II). The transition vesicles move toward the <em>cis</em> Golgi on microtubules. As they do so, their COPII coat is removed and they may fuse together forming larger vesicles; these fuse with the <em>cis</em> Golgi. Sugars are added to proteins in small packets so many glycoproteins have to undergo a large number of sequential steps of glycosylation, each requiring its own enzymes. These steps take place as <strong>shuttle vesicles</strong> carry the proteins from <strong><em>cis</em></strong> to <strong>medial</strong> to the <strong><em>trans</em> Golgi</strong> compartments. At the outer face of the <em>trans</em> Golgi, vesicles pinch off and carry their completed products to their various destinations.</li>
    <li class="lt-bio-3970" style="background-color: unset;"><strong>Mechanism 2:</strong> In addition to the pinching off and fusing of shuttle vesicles, the cisternae of the Golgi actually migrate themselves, that is, the <em>cis</em> Golgi gradually migrates up the stack becoming a medial and finally a <em>trans</em> Golgi (Figure <mjx-container class="MathJax CtxtMenu_Attached_0" jax="SVG" overflow="linebreak" tabindex="0" ctxtmenu_counter="99" style="font-size: 85%; position: relative;"><svg width="4.652ex" height="1.557ex" role="img" focusable="false" viewbox="0 -666 2056 688" aria-hidden="true" style="vertical-align: -0.05ex;"><defs><path id="MJX-100-NCM-N-33" d="M303 353C369 378 431 441 431 526C431 569 410 604 369 631C333 654 292 666 246 666C201 666 162 654 127 631C88 605 68 571 68 528C68 495 90 472 122 472C154 472 176 495 176 527C176 560 157 578 119 580C145 615 186 633 242 633C302 633 332 598 332 527C332 485 324 450 309 421C282 373 245 364 183 364C171 362 165 357 165 348C165 333 172 333 192 333L235 333C310 333 348 280 348 173C348 88 317 14 241 14C176 14 128 36 99 80C134 79 160 105 160 139C160 173 135 198 101 198C62 198 42 178 42 137C42 88 64 49 108 18C147-9 193-22 244-22C301-22 350-3 393 34C436 71 457 117 457 173C457 267 383 332 303 353Z"></path><path id="MJX-100-NCM-N-2E" d="M192 53C192 82 168 106 139 106C110 106 86 82 86 53C86 24 110 0 139 0C168 0 192 24 192 53Z"></path><path id="MJX-100-NCM-N-36" d="M383 504C416 504 432 521 432 555C432 627 378 666 304 666C221 666 155 627 106 548C63 480 42 403 42 316C42 189 65 100 112 47C152 1 198-22 251-22C312-22 362 1 401 47C438 91 457 144 457 205C457 266 439 318 403 361C365 407 316 431 257 431C205 431 165 402 138 346L138 352C138 465 166 561 226 605C252 624 279 633 306 633C342 633 368 623 385 602C351 602 334 583 334 553C334 525 355 504 383 504M344 340C355 317 361 272 361 206C361 141 356 98 345 76C325 35 294 14 251 14C222 14 200 24 184 44C171 60 162 74 158 85C146 116 140 163 140 227C140 255 144 282 151 308C164 355 201 399 256 399C295 399 325 379 344 340Z"></path></defs><g stroke="currentColor" fill="currentColor" stroke-width="0" transform="scale(1,-1)"><g data-mml-node="math" data-latex="\PageIndex{3}" data-semantic-structure="(5 0 1 2 3 4)"><g data-mml-node="TeXAtom" data-mjx-texclass="ORD" data-latex="3}" data-semantic-type="punctuated" data-semantic-role="sequence" data-semantic-annotation="depth:1" data-semantic-id="5" data-semantic-children="0,1,2,3,4" data-semantic-content="1,3" data-semantic-attributes="latex:\PageIndex{3};texclass:ORD" data-semantic-owns="0 1 2 3 4" data-semantic-level-number="0" data-speech-node="true"><g data-mml-node="mn" data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-id="0" data-semantic-parent="5" data-semantic-attributes="latex:3" data-semantic-level-number="1" data-speech-node="true"><use data-c="33" href="#MJX-100-NCM-N-33"></use></g><g data-mml-node="TeXAtom" data-mjx-texclass="ORD" data-latex="{.}" transform="translate(500,0)"><g data-mml-node="mo" data-latex="." data-semantic-type="punctuation" data-semantic-role="fullstop" data-semantic-annotation="nemeth:number;depth:2" data-semantic-id="1" data-semantic-parent="5" data-semantic-attributes="latex:{.};texclass:ORD" data-semantic-operator="punctuated" data-semantic-level-number="1" data-speech-node="true"><use data-c="2E" href="#MJX-100-NCM-N-2E"></use></g></g><g data-mml-node="mn" data-latex="6" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-id="2" data-semantic-parent="5" data-semantic-attributes="latex:6" data-semantic-level-number="1" data-speech-node="true" transform="translate(778,0)"><use data-c="36" href="#MJX-100-NCM-N-36"></use></g><g data-mml-node="TeXAtom" data-mjx-texclass="ORD" data-latex="{.}" transform="translate(1278,0)"><g data-mml-node="mo" data-latex="." data-semantic-type="punctuation" data-semantic-role="fullstop" data-semantic-annotation="nemeth:number;depth:2" data-semantic-id="3" data-semantic-parent="5" data-semantic-attributes="latex:{.};texclass:ORD" data-semantic-operator="punctuated" data-semantic-level-number="1" data-speech-node="true"><use data-c="2E" href="#MJX-100-NCM-N-2E"></use></g></g><g data-mml-node="mn" data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-id="4" data-semantic-parent="5" data-semantic-attributes="latex:3" data-semantic-level-number="1" data-speech-node="true" transform="translate(1556,0)"><use data-c="33" href="#MJX-100-NCM-N-33"></use></g></g></g></g></svg><mjx-assistive-mml unselectable="on" display="inline"><math data-latex="\PageIndex{3}" data-semantic-structure="(5 0 1 2 3 4)"><mrow data-mjx-texclass="ORD" data-latex="3}" data-semantic-type="punctuated" data-semantic-role="sequence" data-semantic-annotation="depth:1" data-semantic-="" data-semantic-children="0,1,2,3,4" data-semantic-content="1,3" data-semantic-attributes="latex:\PageIndex{3};texclass:ORD" data-semantic-owns="0 1 2 3 4" data-semantic-level-number="0" data-speech-node="true"><mn data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:3" data-semantic-level-number="1" data-speech-node="true">3</mn><mrow data-mjx-texclass="ORD" data-latex="{.}"><mo data-latex="." data-semantic-type="punctuation" data-semantic-role="fullstop" data-semantic-annotation="nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:{.};texclass:ORD" data-semantic-operator="punctuated" data-semantic-level-number="1" data-speech-node="true">.</mo></mrow><mn data-latex="6" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:6" data-semantic-level-number="1" data-speech-node="true">6</mn><mrow data-mjx-texclass="ORD" data-latex="{.}"><mo data-latex="." data-semantic-type="punctuation" data-semantic-role="fullstop" data-semantic-annotation="nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:{.};texclass:ORD" data-semantic-operator="punctuated" data-semantic-level-number="1" data-speech-node="true">.</mo></mrow><mn data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:3" data-semantic-level-number="1" data-speech-node="true">3</mn></mrow></math></mjx-assistive-mml></mjx-container>).</li>
</ul>

<figure><img class="internal" alt="Diagram of protein transport from ER through cis, medial, and trans Golgi via COPII and COPI vesicles to plasma membrane or lysosomes" loading="lazy" src="https://bio.libretexts.org/@api/deki/files/5584/Golgi.gif?revision=1" aria-describedby="remedy-description-389cc16da0487e7c" /><span id="remedy-description-389cc16da0487e7c" data-remedy-image-description="389cc16da0487e7c" style="display: block; white-space: pre-line;">Diagram of vesicular protein transport through the Golgi apparatus. At the bottom, the endoplasmic reticulum (ER), drawn as an orange cisterna studded with ribosomes (black dots), contains processing proteins (pink squiggles) and proteins being processed (red bars and green dots). Green COPII-coated transition vesicles bud from the ER and move along microtubules (labeled at left) to the cis Golgi. Proteins then move upward through the stack via shuttle vesicles, indicated by red upward arrows between the cis, medial, and trans Golgi cisternae. Each cisterna contains red bars and green dots (proteins being processed) plus processing proteins. On the right side of the stack, purple COPI-coated vesicles carrying pink squiggle processing proteins move downward in retrograde transport, shown by downward arrows. At the top, three vesicles bud from the trans Golgi and travel upward (black arrows) to the plasma membrane, storage vesicles, or lysosomes. Legend: pink squiggle = processing proteins; red bar &amp; green dot = proteins being processed; green circle = COP II; purple circle = COP I.</span>     <figcaption>Figure <mjx-container class="MathJax CtxtMenu_Attached_0" jax="SVG" overflow="linebreak" tabindex="0" ctxtmenu_counter="100" style="font-size: 85%; position: relative;"><svg width="4.652ex" height="1.557ex" role="img" focusable="false" viewbox="0 -666 2056 688" aria-hidden="true" style="vertical-align: -0.05ex;"><defs><path id="MJX-101-NCM-N-33" d="M303 353C369 378 431 441 431 526C431 569 410 604 369 631C333 654 292 666 246 666C201 666 162 654 127 631C88 605 68 571 68 528C68 495 90 472 122 472C154 472 176 495 176 527C176 560 157 578 119 580C145 615 186 633 242 633C302 633 332 598 332 527C332 485 324 450 309 421C282 373 245 364 183 364C171 362 165 357 165 348C165 333 172 333 192 333L235 333C310 333 348 280 348 173C348 88 317 14 241 14C176 14 128 36 99 80C134 79 160 105 160 139C160 173 135 198 101 198C62 198 42 178 42 137C42 88 64 49 108 18C147-9 193-22 244-22C301-22 350-3 393 34C436 71 457 117 457 173C457 267 383 332 303 353Z"></path><path id="MJX-101-NCM-N-2E" d="M192 53C192 82 168 106 139 106C110 106 86 82 86 53C86 24 110 0 139 0C168 0 192 24 192 53Z"></path><path id="MJX-101-NCM-N-36" d="M383 504C416 504 432 521 432 555C432 627 378 666 304 666C221 666 155 627 106 548C63 480 42 403 42 316C42 189 65 100 112 47C152 1 198-22 251-22C312-22 362 1 401 47C438 91 457 144 457 205C457 266 439 318 403 361C365 407 316 431 257 431C205 431 165 402 138 346L138 352C138 465 166 561 226 605C252 624 279 633 306 633C342 633 368 623 385 602C351 602 334 583 334 553C334 525 355 504 383 504M344 340C355 317 361 272 361 206C361 141 356 98 345 76C325 35 294 14 251 14C222 14 200 24 184 44C171 60 162 74 158 85C146 116 140 163 140 227C140 255 144 282 151 308C164 355 201 399 256 399C295 399 325 379 344 340Z"></path></defs><g stroke="currentColor" fill="currentColor" stroke-width="0" transform="scale(1,-1)"><g data-mml-node="math" data-latex="\PageIndex{3}" data-semantic-structure="(5 0 1 2 3 4)"><g data-mml-node="TeXAtom" data-mjx-texclass="ORD" data-latex="3}" data-semantic-type="punctuated" data-semantic-role="sequence" data-semantic-annotation="depth:1" data-semantic-id="5" data-semantic-children="0,1,2,3,4" data-semantic-content="1,3" data-semantic-attributes="latex:\PageIndex{3};texclass:ORD" data-semantic-owns="0 1 2 3 4" data-semantic-level-number="0" data-speech-node="true"><g data-mml-node="mn" data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-id="0" data-semantic-parent="5" data-semantic-attributes="latex:3" data-semantic-level-number="1" data-speech-node="true"><use data-c="33" href="#MJX-101-NCM-N-33"></use></g><g data-mml-node="TeXAtom" data-mjx-texclass="ORD" data-latex="{.}" transform="translate(500,0)"><g data-mml-node="mo" data-latex="." 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<span id="The_Inbound_Path_(Membrane_Fusion)"></span><span id="The_Inbound_Path_(Membrane_Fusion)-2"></span><h2 style="background-color: unset;" class="lt-bio-3970">The Inbound Path (Membrane Fusion)</h2>

<p class="lt-bio-3970" style="background-color: unset;">The movement of cisternal contents through the stack means that essential processing enzymes are also moving away from their proper site of action. Using a variety of signals, the Golgi separates the products from the processing enzymes that made them and returns the enzymes back to the endoplasmic reticulum. This transport is also done by pinching off vesicles, but the inbound vesicles are coated with <strong>COPI</strong> (<strong>coat protein I</strong>). A vesicle recognize its correct target by involving pairs of <strong>complementary</strong> integral membrane proteins:</p>

<ul>
    <li class="lt-bio-3970" style="background-color: unset;"><strong>v-SNAREs</strong> = &quot;vesicle SNAREs&quot; &mdash; on the vesicle surface</li>
    <li class="lt-bio-3970" style="background-color: unset;"><strong>t-SNAREs</strong> = &quot;target SNAREs&quot; &mdash; on the surface of the target membrane</li>
</ul>

<p class="lt-bio-3970" style="background-color: unset;">v-SNAREs and t-SNAREs bind specifically to each other thanks to the complementary structure of their surface domains. Binding is followed by fusion of the two membranes (Figure <mjx-container class="MathJax CtxtMenu_Attached_0" jax="SVG" overflow="linebreak" tabindex="0" ctxtmenu_counter="101" style="font-size: 85%; position: relative;"><svg width="4.652ex" height="1.581ex" role="img" focusable="false" viewbox="0 -677 2056 699" aria-hidden="true" style="vertical-align: -0.05ex;"><defs><path id="MJX-102-NCM-N-33" d="M303 353C369 378 431 441 431 526C431 569 410 604 369 631C333 654 292 666 246 666C201 666 162 654 127 631C88 605 68 571 68 528C68 495 90 472 122 472C154 472 176 495 176 527C176 560 157 578 119 580C145 615 186 633 242 633C302 633 332 598 332 527C332 485 324 450 309 421C282 373 245 364 183 364C171 362 165 357 165 348C165 333 172 333 192 333L235 333C310 333 348 280 348 173C348 88 317 14 241 14C176 14 128 36 99 80C134 79 160 105 160 139C160 173 135 198 101 198C62 198 42 178 42 137C42 88 64 49 108 18C147-9 193-22 244-22C301-22 350-3 393 34C436 71 457 117 457 173C457 267 383 332 303 353Z"></path><path id="MJX-102-NCM-N-2E" d="M192 53C192 82 168 106 139 106C110 106 86 82 86 53C86 24 110 0 139 0C168 0 192 24 192 53Z"></path><path id="MJX-102-NCM-N-36" d="M383 504C416 504 432 521 432 555C432 627 378 666 304 666C221 666 155 627 106 548C63 480 42 403 42 316C42 189 65 100 112 47C152 1 198-22 251-22C312-22 362 1 401 47C438 91 457 144 457 205C457 266 439 318 403 361C365 407 316 431 257 431C205 431 165 402 138 346L138 352C138 465 166 561 226 605C252 624 279 633 306 633C342 633 368 623 385 602C351 602 334 583 334 553C334 525 355 504 383 504M344 340C355 317 361 272 361 206C361 141 356 98 345 76C325 35 294 14 251 14C222 14 200 24 184 44C171 60 162 74 158 85C146 116 140 163 140 227C140 255 144 282 151 308C164 355 201 399 256 399C295 399 325 379 344 340Z"></path><path id="MJX-102-NCM-N-34" d="M353 677C344 677 336 672 330 663L28 199L28 163L289 163L289 81C289 63 285 51 278 46C271 41 252 39 219 39L194 39L194 0C223 2 269 3 331 3C393 3 439 2 468 0L468 39L443 39C410 39 391 41 384 46C377 51 373 63 373 81L373 163L471 163L471 202L373 202L373 660C373 670 366 677 353 677M295 553L295 202L67 202Z"></path></defs><g stroke="currentColor" fill="currentColor" stroke-width="0" transform="scale(1,-1)"><g data-mml-node="math" data-latex="\PageIndex{4}" data-semantic-structure="(5 0 1 2 3 4)"><g data-mml-node="TeXAtom" data-mjx-texclass="ORD" data-latex="4}" data-semantic-type="punctuated" data-semantic-role="sequence" data-semantic-annotation="depth:1" data-semantic-id="5" data-semantic-children="0,1,2,3,4" data-semantic-content="1,3" data-semantic-attributes="latex:\PageIndex{4};texclass:ORD" data-semantic-owns="0 1 2 3 4" data-semantic-level-number="0" data-speech-node="true"><g data-mml-node="mn" data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-id="0" data-semantic-parent="5" data-semantic-attributes="latex:3" data-semantic-level-number="1" 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data-semantic-parent="5" data-semantic-attributes="latex:4" data-semantic-level-number="1" data-speech-node="true">4</mn></mrow></math></mjx-assistive-mml></mjx-container>).</p>

<figure><img class="internal" alt="Vesicle v-SNARE and receiving-compartment t-SNARE meet at a membrane contact site." loading="lazy" src="https://bio.libretexts.org/@api/deki/files/5585/snares.gif?revision=1" aria-describedby="remedy-description-da7a9bd9bff9b37a" /><span id="remedy-description-da7a9bd9bff9b37a" data-remedy-image-description="da7a9bd9bff9b37a" style="display: block; white-space: pre-line;">A circular vesicle on the left touches the boundary of a receiving compartment on the right. Labels point to v-SNARE at the vesicle membrane and t-SNARE at the target membrane. Red and blue segments meet at the contact point, illustrating the protein interaction that brings the membranes together.</span><figcaption>Figure <mjx-container class="MathJax CtxtMenu_Attached_0" jax="SVG" overflow="linebreak" tabindex="0" ctxtmenu_counter="102" style="font-size: 85%; position: relative;"><svg width="4.652ex" height="1.581ex" role="img" focusable="false" viewbox="0 -677 2056 699" aria-hidden="true" style="vertical-align: -0.05ex;"><defs><path id="MJX-103-NCM-N-33" d="M303 353C369 378 431 441 431 526C431 569 410 604 369 631C333 654 292 666 246 666C201 666 162 654 127 631C88 605 68 571 68 528C68 495 90 472 122 472C154 472 176 495 176 527C176 560 157 578 119 580C145 615 186 633 242 633C302 633 332 598 332 527C332 485 324 450 309 421C282 373 245 364 183 364C171 362 165 357 165 348C165 333 172 333 192 333L235 333C310 333 348 280 348 173C348 88 317 14 241 14C176 14 128 36 99 80C134 79 160 105 160 139C160 173 135 198 101 198C62 198 42 178 42 137C42 88 64 49 108 18C147-9 193-22 244-22C301-22 350-3 393 34C436 71 457 117 457 173C457 267 383 332 303 353Z"></path><path id="MJX-103-NCM-N-2E" d="M192 53C192 82 168 106 139 106C110 106 86 82 86 53C86 24 110 0 139 0C168 0 192 24 192 53Z"></path><path id="MJX-103-NCM-N-36" d="M383 504C416 504 432 521 432 555C432 627 378 666 304 666C221 666 155 627 106 548C63 480 42 403 42 316C42 189 65 100 112 47C152 1 198-22 251-22C312-22 362 1 401 47C438 91 457 144 457 205C457 266 439 318 403 361C365 407 316 431 257 431C205 431 165 402 138 346L138 352C138 465 166 561 226 605C252 624 279 633 306 633C342 633 368 623 385 602C351 602 334 583 334 553C334 525 355 504 383 504M344 340C355 317 361 272 361 206C361 141 356 98 345 76C325 35 294 14 251 14C222 14 200 24 184 44C171 60 162 74 158 85C146 116 140 163 140 227C140 255 144 282 151 308C164 355 201 399 256 399C295 399 325 379 344 340Z"></path><path id="MJX-103-NCM-N-34" d="M353 677C344 677 336 672 330 663L28 199L28 163L289 163L289 81C289 63 285 51 278 46C271 41 252 39 219 39L194 39L194 0C223 2 269 3 331 3C393 3 439 2 468 0L468 39L443 39C410 39 391 41 384 46C377 51 373 63 373 81L373 163L471 163L471 202L373 202L373 660C373 670 366 677 353 677M295 553L295 202L67 202Z"></path></defs><g stroke="currentColor" fill="currentColor" stroke-width="0" transform="scale(1,-1)"><g data-mml-node="math" data-latex="\PageIndex{4}" data-semantic-structure="(5 0 1 2 3 4)"><g data-mml-node="TeXAtom" data-mjx-texclass="ORD" data-latex="4}" data-semantic-type="punctuated" data-semantic-role="sequence" data-semantic-annotation="depth:1" data-semantic-id="5" data-semantic-children="0,1,2,3,4" data-semantic-content="1,3" data-semantic-attributes="latex:\PageIndex{4};texclass:ORD" data-semantic-owns="0 1 2 3 4" data-semantic-level-number="0" data-speech-node="true"><g data-mml-node="mn" data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-id="0" 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href="#MJX-103-NCM-N-34"></use></g></g></g></g></svg><mjx-assistive-mml unselectable="on" display="inline"><math data-latex="\PageIndex{4}" data-semantic-structure="(5 0 1 2 3 4)"><mrow data-mjx-texclass="ORD" data-latex="4}" data-semantic-type="punctuated" data-semantic-role="sequence" data-semantic-annotation="depth:1" data-semantic-="" data-semantic-children="0,1,2,3,4" data-semantic-content="1,3" data-semantic-attributes="latex:\PageIndex{4};texclass:ORD" data-semantic-owns="0 1 2 3 4" data-semantic-level-number="0" data-speech-node="true"><mn data-latex="3" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:2" data-semantic-="" data-semantic-parent="5" data-semantic-attributes="latex:3" data-semantic-level-number="1" data-speech-node="true">3</mn><mrow data-mjx-texclass="ORD" data-latex="{.}"><mo data-latex="." data-semantic-type="punctuation" data-semantic-role="fullstop" 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<div class="box-note"><span id="The_Golgi_is_Not_a_Static_Organelle"></span><span id="The_Golgi_is_Not_a_Static_Organelle-2"></span><h3 class="box-legend"><span class="lt-icon-default">The Golgi is Not a Static Organelle</span></h3>

<p class="lt-bio-3970" style="background-color: unset;">The Golgi breaks up and disappears at the onset of mitosis. By telophase of mitosis, the Golgi reappears. How it is recreated is still uncertain.</p>
</div>

<hr class="autoattribution-divider" /><div class="autoattribution"><p>This page titled <a target="_blank" class="internal mt-self-link" href="/Sandboxes/johnnyphung/biology/03:_The_Cellular_Basis_of_Life/3.06:_Golgi_Apparatus">3.6: Golgi Apparatus</a> is shared under a <a rel="nofollow" href="https://creativecommons.org/licenses/by/3.0" target="_blank">CC BY  3.0 (creativecommons.org)</a>  license and was authored, remixed, and/or curated by <a rel="nofollow" target="_blank" href="http://www.biology-pages.info/">John W. Kimball (biology-pages.info)</a> via <a rel="nofollow" href="https://www.biology-pages.info/" target="_blank">source content (biology-pages.info)</a> that was edited to the style and standards of the LibreTexts platform.</p></div>

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