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4.07:_Photosynthesis_-_Pathway_of_Carbon_Fixation
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<p class="lt-bio-4633" style="background-color: unset;">Photosynthesis is the synthesis of organic molecules using the energy of light. 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data-semantic-children="73" data-semantic-attributes="latex:\ce{6CO2 + 12H2O -> C6H12O6 + 6H2O + 6O2}" data-semantic-owns="73" data-semantic-level-number="0" data-speech-node="true"><mlabeledtr data-semantic-type="line" data-semantic-role="multiline" data-semantic-annotation="depth:2" data-semantic-="" data-semantic-children="69" data-semantic-content="4" data-semantic-parent="74" data-semantic-owns="69 4" data-semantic-level-number="1" data-speech-node="true"><mtd data-semantic-type="cell" data-semantic-role="label" data-semantic-annotation="depth:3" data-semantic-id="4" data-semantic-children="3" data-semantic-parent="73" data-semantic-owns="3" data-semantic-level-number="2" data-speech-node="true"><mrow data-semantic-added="true" data-semantic-type="fenced" data-semantic-role="leftright" data-semantic-annotation="fences:paren;depth:4" data-semantic-="" data-semantic-children="1" data-semantic-content="0,2" data-semantic-parent="4" data-semantic-owns="0 1 2" data-semantic-level-number="3" data-speech-node="true"><mtext data-latex="\text{(}" data-semantic-type="fence" data-semantic-role="open" data-semantic-annotation="general:text;depth:5" data-semantic-="" data-semantic-parent="3" data-semantic-attributes="latex:\text{(}" data-semantic-operator="fenced" data-semantic-level-number="4" data-speech-node="true">(</mtext><mtext data-latex="\text{4.7.1}" data-semantic-type="text" data-semantic-role="annotation" data-semantic-font="normal" data-semantic-annotation="clearspeak:unit;depth:5" data-semantic-="" data-semantic-parent="3" data-semantic-attributes="latex:\text{4.7.1}" data-semantic-level-number="4" data-speech-node="true">4.7.1</mtext><mtext data-latex="\text{)}" data-semantic-type="fence" data-semantic-role="close" data-semantic-annotation="general:text;depth:5" data-semantic-="" data-semantic-parent="3" data-semantic-attributes="latex:\text{)}" data-semantic-operator="fenced" data-semantic-level-number="4" data-speech-node="true">)</mtext></mrow></mtd><mtd><mrow data-mjx-texclass="ORD" data-latex="{6\,\mathrm{CO}{\vphantom{A}}_{\smash[t]{2}}{}+{}12\,\mathrm{H}{\vphantom{A}}_{\smash[t]{2}}\mathrm{O}{}\mathrel{\mhchemlongrightarrow}{}\mathrm{C}{\vphantom{A}}_{\smash[t]{6}}\mathrm{H}{\vphantom{A}}_{\smash[t]{12}}\mathrm{O}{\vphantom{A}}_{\smash[t]{6}}{}+{}6\,\mathrm{H}{\vphantom{A}}_{\smash[t]{2}}\mathrm{O}{}+{}6\,\mathrm{O}{\vphantom{A}}_{\smash[t]{2}}}"><mrow data-semantic-added="true" data-semantic-type="infixop" data-semantic-role="addition" data-semantic-annotation="depth:3" data-semantic-="" data-semantic-children="66,68,71" data-semantic-content="32,40" data-semantic-parent="73" data-semantic-attributes="latex:{6\,\mathrm{CO}{\vphantom{A}}_{\smash[t]{2}}{}+{}12\,\mathrm{H}{\vphantom{A}}_{\smash[t]{2}}\mathrm{O}{}\mathrel{\mhchemlongrightarrow}{}\mathrm{C}{\vphantom{A}}_{\smash[t]{6}}\mathrm{H}{\vphantom{A}}_{\smash[t]{12}}\mathrm{O}{\vphantom{A}}_{\smash[t]{6}}{}+{}6\,\mathrm{H}{\vphantom{A}}_{\smash[t]{2}}\mathrm{O}{}+{}6\,\mathrm{O}{\vphantom{A}}_{\smash[t]{2}}};texclass:ORD" data-semantic-owns="66 32 68 40 71" data-semantic-level-number="2" data-speech-node="true"><mrow data-semantic-added="true" data-semantic-type="infixop" data-semantic-role="unknown" data-semantic-annotation="depth:4" data-semantic-="" data-semantic-children="63,65" data-semantic-content="19" data-semantic-parent="69" data-semantic-owns="63 19 65" data-semantic-level-number="3" data-speech-node="true"><mrow data-semantic-added="true" data-semantic-type="infixop" data-semantic-role="addition" data-semantic-annotation="depth:5" data-semantic-="" data-semantic-children="60,62" data-semantic-content="11" data-semantic-parent="66" data-semantic-owns="60 11 62" data-semantic-level-number="4" data-speech-node="true"><mrow data-semantic-added="true" data-semantic-type="infixop" data-semantic-role="implicit" data-semantic-annotation="clearspeak:unit;depth:6" data-semantic-="" data-semantic-children="5,10" data-semantic-content="59" data-semantic-parent="63" data-semantic-owns="5 59 10" data-semantic-level-number="5" data-speech-node="true"><mn data-latex="6" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:7" data-semantic-="" data-semantic-parent="60" data-semantic-attributes="latex:6" data-semantic-level-number="6" data-speech-node="true">6</mn><mspace width="0.167em" data-latex="\," data-semantic-type="operator" data-semantic-role="space" data-semantic-annotation="depth:7" data-semantic-="" data-semantic-parent="60" data-semantic-operator="infixop,⁢" data-semantic-level-number="6" data-speech-node="true"></mspace><mrow data-mjx-texclass="ORD" data-latex="\mathrm{CO}"><mrow data-semantic-added="true" data-semantic-type="subscript" data-semantic-role="identifier" data-semantic-annotation="collapsed:identifier;depth:7" data-semantic-="" data-semantic-children="7,9" data-semantic-parent="60" data-semantic-attributes="latex:{\vphantom{A}}_{\smash[t]{2}}" data-semantic-owns="7 9" data-semantic-level-number="6" data-speech-node="true"><mi data-mjx-auto-op="false" data-latex="CO" data-semantic-type="identifier" data-semantic-role="unknown" data-semantic-font="normal" data-semantic-annotation="depth:8" data-semantic-="" data-semantic-parent="10" data-semantic-attributes="latex:\mathrm{CO};texclass:ORD" data-semantic-level-number="7" data-speech-node="true">CO</mi></mrow></mrow></mrow><msub data-latex="{\vphantom{A}}_{\smash[t]{2}}"><mrow data-mjx-texclass="ORD" data-latex="{\vphantom{A}}"><mrow data-mjx-texclass="ORD" data-latex="\vphantom{A}"><mpadded width="0"><mphantom><mi data-latex="A">A</mi></mphantom></mpadded></mrow></mrow><mrow data-mjx-texclass="ORD" data-latex="{\smash[t]{2}}"><mrow data-mjx-texclass="ORD" data-latex="\smash[t]{2}"><mpadded height="0"><mn data-latex="2" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;depth:8" data-semantic-="" data-semantic-parent="10" data-semantic-attributes="latex:{\smash[t]{2}};texclass:ORD" data-semantic-level-number="7" data-speech-node="true">2</mn></mpadded></mrow></mrow></msub><mrow data-mjx-texclass="ORD" data-latex="{}"></mrow><mo data-latex="+" data-semantic-type="operator" data-semantic-role="addition" data-semantic-annotation="depth:6" data-semantic-="" data-semantic-parent="63" data-semantic-attributes="latex:+" data-semantic-operator="infixop,+" data-semantic-level-number="5" data-speech-node="true">+</mo><mrow data-mjx-texclass="ORD" data-latex="{}"></mrow><mrow data-semantic-added="true" data-semantic-type="infixop" data-semantic-role="implicit" data-semantic-annotation="clearspeak:unit;depth:6" data-semantic-="" data-semantic-children="12,58" data-semantic-content="61" data-semantic-parent="63" data-semantic-owns="12 61 58" data-semantic-level-number="5" data-speech-node="true"><mn data-latex="12" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;depth:7" data-semantic-="" data-semantic-parent="62" data-semantic-attributes="latex:12" data-semantic-level-number="6" data-speech-node="true">12</mn><mspace width="0.167em" data-latex="\,"></mspace><mo data-semantic-type="operator" data-semantic-role="multiplication" data-semantic-annotation="depth:7" data-semantic-="" data-semantic-parent="62" data-semantic-added="true" data-semantic-operator="infixop,⁢" data-semantic-level-number="6" data-speech-node="true">⁢</mo><mrow data-semantic-added="true" data-semantic-type="appl" data-semantic-role="simple function" data-semantic-annotation="depth:7" data-semantic-="" data-semantic-children="14,56" data-semantic-content="57,14" data-semantic-parent="62" data-semantic-owns="14 57 56" data-semantic-level-number="6" data-speech-node="true"><mrow data-mjx-texclass="ORD" data-latex="\mathrm{H}"><mi mathvariant="normal" data-latex="H" data-semantic-type="identifier" data-semantic-role="simple function" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;nemeth:number;depth:8" data-semantic-="" data-semantic-parent="58" data-semantic-attributes="latex:\mathrm{H};texclass:ORD" data-semantic-operator="appl" data-semantic-level-number="7" data-speech-node="true">H</mi></mrow><mo data-semantic-type="punctuation" data-semantic-role="application" data-semantic-annotation="nemeth:number;depth:8" data-semantic-="" data-semantic-parent="58" data-semantic-added="true" data-semantic-operator="appl" data-semantic-level-number="7" data-speech-node="true">⁡</mo><mrow data-semantic-added="true" data-semantic-type="infixop" data-semantic-role="implicit" data-semantic-annotation="clearspeak:unit;depth:8" data-semantic-="" data-semantic-children="17,18" data-semantic-content="55" data-semantic-parent="58" data-semantic-owns="17 55 18" data-semantic-level-number="7" data-speech-node="true"><msub data-latex="{\vphantom{A}}_{\smash[t]{2}}" data-semantic-type="subscript" data-semantic-role="unknown" data-semantic-annotation="depth:9" data-semantic-="" data-semantic-children="15,16" data-semantic-parent="56" data-semantic-attributes="latex:{\vphantom{A}}_{\smash[t]{2}}" data-semantic-owns="15 16" data-semantic-level-number="8" data-speech-node="true"><mrow data-mjx-texclass="ORD" data-latex="{\vphantom{A}}"><mrow data-mjx-texclass="ORD" data-latex="\vphantom{A}" data-semantic-type="empty" data-semantic-role="unknown" data-semantic-annotation="nemeth:number;depth:10" data-semantic-="" data-semantic-parent="17" data-semantic-attributes="latex:{\vphantom{A}};texclass:ORD" data-semantic-level-number="9" data-speech-node="true"><mpadded width="0"><mphantom><mi data-latex="A">A</mi></mphantom></mpadded></mrow></mrow><mrow data-mjx-texclass="ORD" data-latex="{\smash[t]{2}}"><mrow data-mjx-texclass="ORD" data-latex="\smash[t]{2}"><mpadded height="0"><mn data-latex="2" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;depth:10" data-semantic-="" data-semantic-parent="17" data-semantic-attributes="latex:{\smash[t]{2}};texclass:ORD" data-semantic-level-number="9" data-speech-node="true">2</mn></mpadded></mrow></mrow></msub><mo data-semantic-type="operator" data-semantic-role="multiplication" data-semantic-annotation="depth:9" data-semantic-="" data-semantic-parent="56" data-semantic-added="true" data-semantic-operator="infixop,⁢" data-semantic-level-number="8" data-speech-node="true">⁢</mo><mrow data-mjx-texclass="ORD" data-latex="\mathrm{O}"><mi mathvariant="normal" data-latex="O" data-semantic-type="identifier" data-semantic-role="latinletter" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;depth:9" data-semantic-="" data-semantic-parent="56" data-semantic-attributes="latex:\mathrm{O};texclass:ORD" data-semantic-level-number="8" data-speech-node="true">O</mi></mrow></mrow></mrow></mrow></mrow><mrow data-mjx-texclass="ORD" data-latex="{}"></mrow><mrow data-mjx-texclass="REL" data-latex="\mathrel{\mhchemlongrightarrow}"><mo data-mjx-variant="-mhchem" data-mjx-texclass="REL" stretchy="true" data-latex="\mhchemlongrightarrow" data-semantic-type="operator" data-semantic-role="unknown" data-semantic-annotation="depth:5" data-semantic-="" data-semantic-parent="66" data-semantic-attributes="latex:\mathrel{\mhchemlongrightarrow};texclass:REL" data-semantic-operator="infixop," data-semantic-level-number="4" data-speech-node="true"></mo></mrow><mrow data-mjx-texclass="ORD" data-latex="{}"></mrow><mrow data-semantic-added="true" data-semantic-type="infixop" data-semantic-role="implicit" data-semantic-annotation="clearspeak:unit;depth:5" data-semantic-="" data-semantic-children="23,54" data-semantic-content="64" data-semantic-parent="66" data-semantic-owns="23 64 54" data-semantic-level-number="4" data-speech-node="true"><mrow data-mjx-texclass="ORD" data-latex="\mathrm{C}"><mrow data-semantic-added="true" data-semantic-type="subscript" data-semantic-role="identifier" data-semantic-annotation="collapsed:identifier;depth:6" data-semantic-="" data-semantic-children="20,22" data-semantic-parent="65" data-semantic-attributes="latex:{\vphantom{A}}_{\smash[t]{6}}" data-semantic-owns="20 22" data-semantic-level-number="5" data-speech-node="true"><mi mathvariant="normal" data-latex="C" data-semantic-type="identifier" data-semantic-role="latinletter" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;depth:7" data-semantic-="" data-semantic-parent="23" data-semantic-attributes="latex:\mathrm{C};texclass:ORD" data-semantic-level-number="6" data-speech-node="true">C</mi></mrow></mrow><msub data-latex="{\vphantom{A}}_{\smash[t]{6}}"><mrow data-mjx-texclass="ORD" data-latex="{\vphantom{A}}"><mrow data-mjx-texclass="ORD" data-latex="\vphantom{A}"><mpadded width="0"><mphantom><mi data-latex="A">A</mi></mphantom></mpadded></mrow></mrow><mrow data-mjx-texclass="ORD" data-latex="{\smash[t]{6}}"><mrow data-mjx-texclass="ORD" data-latex="\smash[t]{6}"><mpadded height="0"><mn data-latex="6" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;depth:7" data-semantic-="" data-semantic-parent="23" data-semantic-attributes="latex:{\smash[t]{6}};texclass:ORD" data-semantic-level-number="6" data-speech-node="true">6</mn></mpadded></mrow></mrow></msub><mo data-semantic-type="operator" data-semantic-role="multiplication" 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data-mjx-texclass="ORD" data-latex="{\vphantom{A}}"><mrow data-mjx-texclass="ORD" data-latex="\vphantom{A}" data-semantic-type="empty" data-semantic-role="unknown" data-semantic-annotation="nemeth:number;depth:9" data-semantic-="" data-semantic-parent="27" data-semantic-attributes="latex:{\vphantom{A}};texclass:ORD" data-semantic-level-number="8" data-speech-node="true"><mpadded width="0"><mphantom><mi data-latex="A">A</mi></mphantom></mpadded></mrow></mrow><mrow data-mjx-texclass="ORD" data-latex="{\smash[t]{12}}"><mrow data-mjx-texclass="ORD" data-latex="\smash[t]{12}"><mpadded height="0"><mn data-latex="12" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;depth:9" data-semantic-="" data-semantic-parent="27" data-semantic-attributes="latex:{\smash[t]{12}};texclass:ORD" data-semantic-level-number="8" data-speech-node="true">12</mn></mpadded></mrow></mrow></msub><mo data-semantic-type="operator" data-semantic-role="multiplication" data-semantic-annotation="depth:8" data-semantic-="" data-semantic-parent="52" data-semantic-added="true" data-semantic-operator="infixop,⁢" data-semantic-level-number="7" data-speech-node="true">⁢</mo><mrow data-mjx-texclass="ORD" data-latex="\mathrm{O}"><mrow data-semantic-added="true" data-semantic-type="subscript" data-semantic-role="identifier" data-semantic-annotation="collapsed:identifier;depth:8" data-semantic-="" data-semantic-children="28,30" data-semantic-parent="52" data-semantic-attributes="latex:{\vphantom{A}}_{\smash[t]{6}}" data-semantic-owns="28 30" data-semantic-level-number="7" data-speech-node="true"><mi mathvariant="normal" data-latex="O" data-semantic-type="identifier" data-semantic-role="latinletter" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;depth:9" data-semantic-="" data-semantic-parent="31" data-semantic-attributes="latex:\mathrm{O};texclass:ORD" data-semantic-level-number="8" 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data-latex="\mathrm{O}"><mrow data-semantic-added="true" data-semantic-type="subscript" data-semantic-role="identifier" data-semantic-annotation="collapsed:identifier;depth:5" data-semantic-="" data-semantic-children="43,45" data-semantic-parent="71" data-semantic-attributes="latex:{\vphantom{A}}_{\smash[t]{2}}" data-semantic-owns="43 45" data-semantic-level-number="4" data-speech-node="true"><mi mathvariant="normal" data-latex="O" data-semantic-type="identifier" data-semantic-role="latinletter" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;depth:6" data-semantic-="" data-semantic-parent="46" data-semantic-attributes="latex:\mathrm{O};texclass:ORD" data-semantic-level-number="5" data-speech-node="true">O</mi></mrow></mrow></mrow></mrow><msub data-latex="{\vphantom{A}}_{\smash[t]{2}}"><mrow data-mjx-texclass="ORD" data-latex="{\vphantom{A}}"><mrow data-mjx-texclass="ORD" data-latex="\vphantom{A}"><mpadded width="0"><mphantom><mi 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data-semantic-structure="(3 0 2)"><mrow data-mjx-texclass="ORD" data-latex="{\mathrm{CO}{\vphantom{A}}_{\smash[t]{2}}}" data-semantic-type="subscript" data-semantic-role="identifier" data-semantic-annotation="collapsed:identifier;depth:1" data-semantic-="" data-semantic-children="0,2" data-semantic-attributes="latex:\ce{CO2};texclass:ORD" data-semantic-owns="0 2" data-semantic-level-number="0" data-speech-node="true"><mrow data-mjx-texclass="ORD" data-latex="\mathrm{CO}"><mi data-mjx-auto-op="false" data-latex="CO" data-semantic-type="identifier" data-semantic-role="unknown" data-semantic-font="normal" data-semantic-annotation="nemeth:number;depth:2" data-semantic-="" data-semantic-parent="3" data-semantic-attributes="latex:\mathrm{CO};texclass:ORD" data-semantic-level-number="1" data-speech-node="true">CO</mi></mrow><msub data-latex="{\vphantom{A}}_{\smash[t]{2}}"><mrow data-mjx-texclass="ORD" data-latex="{\vphantom{A}}"><mrow data-mjx-texclass="ORD" data-latex="\vphantom{A}"><mpadded width="0"><mphantom><mi data-latex="A">A</mi></mphantom></mpadded></mrow></mrow><mrow data-mjx-texclass="ORD" data-latex="{\smash[t]{2}}"><mrow data-mjx-texclass="ORD" data-latex="\smash[t]{2}"><mpadded height="0"><mn data-latex="2" data-semantic-type="number" data-semantic-role="integer" data-semantic-font="normal" data-semantic-annotation="clearspeak:simple;depth:2" data-semantic-="" data-semantic-parent="3" data-semantic-attributes="latex:{\smash[t]{2}};texclass:ORD" data-semantic-level-number="1" data-speech-node="true">2</mn></mpadded></mrow></mrow></msub></mrow></math></mjx-assistive-mml></mjx-container>) to organic molecules.</p> <span id="The_Steps_that_lead_to_Photosynthesis"></span><span id="The_Steps_that_lead_to_Photosynthesis"></span><h2 style="background-color: unset;" class="lt-bio-4633">The Steps that lead to Photosynthesis</h2> <figure><img class="internal" alt="Flowchart of the Calvin Cycle showing chemical reactions and compounds, including ATP, NADPH, and RuBP, involved in photosynthesis. Highlighted steps include conversion of 3-PGA to G3P." loading="lazy" src="https://bio.libretexts.org/@api/deki/files/6297/CalvinCycle.png?revision=1" width="531" height="393" /><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="1.581ex" role="img" focusable="false" viewbox="0 -677 2056 699" aria-hidden="true" style="vertical-align: -0.05ex;"><defs><path id="MJX-101-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><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 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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:1" data-semantic-level-number="1" data-speech-node="true" transform="translate(1556,0)"><use data-c="31" href="#MJX-101-NCM-N-31"></use></g></g></g></g></svg><mjx-assistive-mml unselectable="on" 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="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 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="7" 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:7" data-semantic-level-number="1" data-speech-node="true">7</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>: The steps in the fixation of carbon dioxide during photosynthesis</figcaption></figure> <ul> <li class="lt-bio-4633" style="background-color: unset;">CO<sub><font size="2">2</font></sub> combines with the phosphorylated 5-carbon sugar <strong>ribulose bisphosphate</strong>.</li> <li class="lt-bio-4633" style="background-color: unset;">This reaction is catalyzed by the enzyme <strong>ribulose bisphosphate carboxylase oxygenase</strong> (<strong>RUBISCO</strong>)(an enzyme which can fairly claim to be the most abundant protein on earth).</li> <li class="lt-bio-4633" style="background-color: unset;">The resulting 6-carbon compound breaks down into two molecules of <strong>3-phosphoglyceric acid</strong> (<strong>PGA</strong>).</li> <li class="lt-bio-4633" style="background-color: unset;">The PGA molecules are further phosphorylated (by <strong>ATP</strong>) and are reduced (by <strong>NADPH</strong>) to form <strong>phosphoglyceraldehyde</strong> (<strong>PGAL</strong>).</li> <li class="lt-bio-4633" style="background-color: unset;">Phosphoglyceraldehyde serves as the starting material for the synthesis of <strong>glucose</strong> and <strong>fructose</strong>.</li> <li class="lt-bio-4633" style="background-color: unset;">Glucose and fructose make the disaccharide <strong>sucrose</strong>, which travels in solution to other parts of the plant (e.g., fruit, roots).</li> <li class="lt-bio-4633" style="background-color: unset;">Glucose is also the monomer used in the synthesis of the <strong>polysaccharides</strong> <strong>starch</strong> and <strong>cellulose</strong>.</li> </ul> <p class="lt-bio-4633" style="background-color: unset;">The Figure 4.7.1 shows the steps in the fixation of carbon dioxide during photosynthesis. All of these reactions occur in the <strong>stroma</strong> of the chloroplast. These steps were worked out by Melvin Calvin and his colleagues at the University of California and, for this reason, are named the <strong>Calvin cycle</strong>.</p> <span id="Calvin's_Experiment"></span><span id="Calvin's_Experiment"></span><h2 style="background-color: unset;" class="lt-bio-4633">Calvin's Experiment</h2> <figure><img class="internal" alt="A black and white image of a laboratory setup with mirrors, glass equipment, and a round-bottom flask. Various light sources and optical devices are positioned around the experimental apparatus." loading="lazy" src="https://bio.libretexts.org/@api/deki/files/6298/Bassham1.jpg?revision=1" /><figcaption>Figure 4.7.2 Apparatus used for Calvin's experiment courtesy of Dr. James A. Bassham</figcaption></figure> <p class="lt-bio-4633" style="background-color: unset;">The experimental apparatus is shown above. After various intervals of illumination, a suspension of unicellular algae is inactivated and the contents of the cells extracted. The compounds in a drop of the extract are then separated by paper chromatography.</p> <p class="lt-bio-4633" style="background-color: unset;">The identity of each substance may be determined simply by comparing its position with the positions occupied by known substances under the same conditions. Or, a fragment containing the spot can be cut from the sheet and chemically analyzed.</p> <p class="lt-bio-4633" style="background-color: unset;">To determine which, if any, of the substances separated on the chromatogram are radioactive, a sheet of X-ray film is placed next to the chromatogram. If dark spots appear on the film (because of radiation emitted by the <sup><small><font size="1">14</font></small></sup>C atoms), their position can be correlated with the positions of the chemicals in the chromatogram. Using this technique of <strong>autoradiography</strong>, Calvin found that <sup><small><font size="1">14</font></small></sup>C turned up in glucose molecules within 30 seconds after the start of photosynthesis. When he permitted photosynthesis to proceed for only 5 seconds, however, the radioactivity was concentrated in several other, smaller, molecules.</p> <figure><img class="internal" alt="A scientific image with labeled spots showing substances: glucose, glycogen, trehalose, ATP, fumaric, pyruvic, fumaric-P, triose-P, glucose-6-P, and gluconic. Text indicates exposed 10 sec." loading="lazy" src="https://bio.libretexts.org/@api/deki/files/6299/Bassham2.jpg?revision=1" /> <img class="internal" alt="A blurry image showing various labeled chemical compounds in red text on a gradient background. The labels include Glucose, Pyruvate, and others. Some labels are partially obscured." loading="lazy" src="https://bio.libretexts.org/@api/deki/files/6300/Bassham3.jpg?revision=1" /><figcaption>Figure 4.7.3 Chromatogram of Calvin's experiment courtesy of Dr. James A. Bassham</figcaption></figure> <p class="lt-bio-4633" style="background-color: unset;">The dark spots show the radioactive compounds produced after 10 secs (left) and 2 minutes (right) of photosynthesis by the green alga Scenedesmus. The alga was supplied with carbon dioxide labeled with <sup><small><font size="1">14</font></small></sup>C, a radioactive isotope of carbon. At 10 seconds, most of the radioactivity is found in 3-phosphoglyceric acid ("P-Glyceric"). At 2 minutes, phosphorylated 6-carbon sugars (glucose and fructose) have been synthesized as well as a number of amino acids. 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