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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;">α-MSH produced by neurons in the arcuate nucleus has important roles in the regulation of appetite and sexual behavior, while α-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;">β-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 depicting the breakdown of Proopiomelanocortin (POMC) into peptides: γ-MSH, ACTH, α-MSH, CLIP, β-Lipotropin, β-MSH, γ-Lipotropin, and β-Endorphin." loading="lazy" width="1280" height="717" src="https://bio.libretexts.org/@api/deki/files/5588/POMC.png?revision=1&size=bestfit&width=602&height=336" /><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 18C147-9 193-22 244-22C301-22 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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 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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="Black and white electron microscope image showing cellular structures with irregular shapes, scattered vesicles, and stacked membranes. A red box highlights two areas with layered formations." loading="lazy" src="https://bio.libretexts.org/@api/deki/files/5583/Golgi.jpg?revision=1" /><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 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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)"></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 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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="Flowchart showing MAPK/ERK pathway: RTK activates Grb2, leading to SOS, Ras, B-Raf, MEK, ERK activation. ERK affects cell cycle. Includes positive and negative feedback loops, and regulatory proteins." loading="lazy" src="https://bio.libretexts.org/@api/deki/files/5584/Golgi.gif?revision=1" /> <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 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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></figcaption></figure> <span id="The_Inbound_Path_(Membrane_Fusion)"></span><span id="The_Inbound_Path_(Membrane_Fusion)"></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> = "vesicle SNAREs" — on the vesicle surface</li> <li class="lt-bio-3970" style="background-color: unset;"><strong>t-SNAREs</strong> = "target SNAREs" — 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 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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" 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="4" 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:4" data-semantic-level-number="1" data-speech-node="true">4</mn></mrow></math></mjx-assistive-mml></mjx-container></figcaption></figure> <div class="box-note"><span id="The_Golgi_is_Not_a_Static_Organelle"></span><span id="The_Golgi_is_Not_a_Static_Organelle"></span><h5 class="box-legend"><span class="lt-icon-default">The Golgi is Not a Static Organelle</span></h5> <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. 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