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4.09:_Photosynthesis_-_Dicovering_the_Secrets
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<p class="lt-bio-4635">This chapter talks about various scientists and their path towards discovering photosynthesis.</p> <span id="van_Helmont"></span><span id="van_Helmont"></span><h2 class="lt-bio-4635">van Helmont</h2> <p class="lt-bio-4635">Perhaps the first experiment designed to explore the nature of photosynthesis was that reported by the Dutch physician van Helmont in 1648. Some years earlier, van Helmont had placed in a large pot exactly 200 pounds (91 kg) of soil that had been thoroughly dried in an oven. Then he moistened the soil with rain water and planted a 5-pound (2.3 kg) willow shoot in it. He then placed the pot in the ground and covered its rim with a perforated iron plate. The perforations allowed water and air to reach the soil but lessened the chance that dirt or other debris would be blown into the pot from the outside.</p> <p class="lt-bio-4635">For five years, van Helmont kept his plant watered with rain water or distilled water. At the end of that time, he carefully removed the young tree and found that it had gained 164 pound, 3 ounces (74.5 kg). (This figure did not include the weight of the leaves that had been shed during the previous four autumns.) He then redried the soil and found that it weighed only 2 ounces (57 g) less that the original 200 pounds (91 kg). Faced with these experimental facts, van Helmont theorized that the increase in weight of the willow arose from the water alone. He did not consider the possibility that gases in the air might be involved.</p> <span id="Joseph_Priestley"></span><span id="Joseph_Priestley"></span><h2 class="lt-bio-4635">Joseph Priestley</h2> <p class="lt-bio-4635">The first evidence that gases participate in photosynthesis was reported by Joseph Priestley in 1772. He knew that if a burning candle is placed in a sealed chamber, the candle soon goes out. If a mouse is then placed in the chamber, it soon suffocates because the process of combustion has used up all the oxygen in the air — the gas on which animal respiration depends. However, Priestley discovered that if a plant is placed in an atmosphere lacking oxygen, it soon replenishes the oxygen, and a mouse can survive in the resulting mixture. Priestley thought (erroneously) that it was simply the growth of the plant that accounted for this.</p> <span id="Ingen-Housz"></span><span id="Ingen-Housz"></span><h2 class="lt-bio-4635">Ingen-Housz</h2> <p class="lt-bio-4635">It was another Dutch physician, Ingen-Housz, who discovered in 1778 that the effect observed by Priestley occurred only when the plant was illuminated. A plant kept in the dark in a sealed chamber consumes oxygen just as a mouse (or candle) does.</p> <p class="lt-bio-4635">Ingen-Housz also demonstrated that only green parts of plants liberated oxygen during photosynthesis. Nongreen plant structure, such as woody stems, roots, flowers, and fruits actually consume oxygen in the process of respiration. We now know that this is because photosynthesis can go on only in the presence of the green pigment chlorophyll.</p> <span id="Jean_Senebier"></span><span id="Jean_Senebier"></span><h2 class="lt-bio-4635">Jean Senebier</h2> <p class="lt-bio-4635">The growth of plants is accompanied by an increase in their carbon content. A Swiss minister, Jean Senebier, discovered that the source of this carbon is carbon dioxide and that the release of oxygen during photosynthesis accompanies the uptake of carbon dioxide. Senebier concluded (erroneously as it turned out) that in photosynthesis carbon dioxide is decomposed, with the carbon becoming incorporated in the organic matter of the plant and the oxygen being released.</p> <p class="mt-indent-3 lt-bio-4635">CO<sub><small><font size="1">2</font></small></sub> + H<sub><small><font size="1">2</font></small></sub>O → (CH<sub><small><font size="1">2</font></small></sub>O) + O<sub><small><font size="1">2</font></small></sub></p> <p class="lt-bio-4635">(The parentheses around the CH<sub><small><font size="1">2</font></small></sub>O signify that no specific molecule is being indicated but, instead, the ratio of atoms in some carbohydrate, e.g., glucose, C<sub><small><font size="1">6</font></small></sub>H<sub><small><font size="1">12</font></small></sub>O<sub><small><font size="1">6</font></small></sub>.) The equation also indicates that the ratio of carbon dioxide consumed to oxygen release is 1:1, a finding that was carefully demonstrated in the years following Senebier's work. Using glucose as the carbohydrate product, we can write the equation for photosynthesis as</p> <p class="mt-indent-3 lt-bio-4635">6CO<sub><small><font size="1">2</font></small></sub> + 6H<sub><small><font size="1">2</font></small></sub>O → C<sub><small><font size="1">6</font></small></sub>H<sub><small><font size="1">12</font></small></sub>O<sub><small><font size="1">6</font></small></sub> + 6O<sub><small><font size="1">2</font></small></sub></p> <span id="F._F._Blackman"></span><span id="F._F._Blackman"></span><h2 class="lt-bio-4635">F. F. Blackman</h2> <figure><img style="width: 234px; height: 225px;" class="internal" alt="Diagram of a photosynthesis experiment with a test tube containing aquatic plants, submerged in water. An arrow labeled light points to the setup, indicating exposure to light." loading="lazy" width="234px" height="225px" src="https://bio.libretexts.org/@api/deki/files/6325/Elodea.jpg?revision=1&size=bestfit&width=234&height=225" /> <img style="width: 282px; height: 260px;" class="internal" alt="Graph showing the rate of photosynthesis vs. light intensity. Two lines represent 20°C and 30°C with phases: light-limited and dark-limited. Higher photosynthesis rate at 30°C with excess CO₂." loading="lazy" width="282px" height="260px" src="https://bio.libretexts.org/@api/deki/files/6326/RateLimits.jpg?revision=1&size=bestfit&width=282&height=260" /><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="1.581ex" role="img" focusable="false" viewbox="0 -677 2056 699" aria-hidden="true" style="vertical-align: -0.05ex;"><defs><path id="MJX-97-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-97-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 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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="9" 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:9" data-semantic-level-number="1" data-speech-node="true">9</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>: (left) Elodea experiment by Blackman (right) Blackman experiment graphic interpretation</figcaption></figure> <p class="lt-bio-4635">The above equation shows the relationship between the substances used in and produced by the process. It tells us nothing about the intermediate steps. That photosynthesis does involve at least two quite distinct processes became apparent from the experiments of the British plant physiologist F. F. Blackman. His results can easily be duplicated by using the setup in Figure 4.9.1. The green water plant <em>Elodea</em> (available wherever aquarium supplies are sold) is the test organism. When a sprig is placed upside down in a dilute solution of NaHCO<sub><small><font size="1">3</font></small></sub> (which serves as a source of CO<sub><small><font size="1">2</font></small></sub>) and illuminated with a flood lamp, oxygen bubbles are soon given off from the cut portion of the stem. One then counts the number of bubbles given off in a fixed interval of time at each of several light intensities. Plotting these data produces a graph like the one in Figure 4.9.2.</p> <p class="lt-bio-4635">Since the rate of photosynthesis does not continue to increase indefinitely with increased illumination, Blackman concluded that at least two distinct processes are involved: one, a reaction that requires light and the other, a reaction that does not. This latter is called a "dark" reaction although it <strong>can</strong> go on in the light. Blackman theorized that at moderate light intensities, the "light" reaction limits or "paces" the entire process. In other words, at these intensities the dark reaction is capable of handling all the intermediate substances produced by the light reaction. With increasing light intensities, however, a point is eventually reached when the dark reaction is working at maximum capacity. Any further illumination is ineffective, and the process reaches a steady rate.</p> <p class="lt-bio-4635">This interpretation is strengthened by repeating the experiment as a somewhat higher temperature. Most chemical reactions proceed more rapidly at higher temperatures (up to a point). At 35°C, the rate of photosynthesis does not level off until greater light intensities are present. This suggest that the dark reaction is now working faster. The fact that at low light intensities the rate of photosynthesis is no greater at 35°C than at 20°C also supports the idea that it is a light reaction that is limiting the process in this range. Light reactions depend, not on temperature, but simply on the intensity of illumination.</p> <p class="lt-bio-4635">The increased rate of photosynthesis with increased temperature does not occur if the supply of CO<sub><small><font size="1">2</font></small></sub> is limited. As the figure shows, the overall rate of photosynthesis reaches a steady value at lower light intensities if the amount of CO<sub><small><font size="1">2</font></small></sub> available is limited. Thus CO<sub><small><font size="1">2</font></small></sub> concentration must be added as a third factor regulating the rate at which photosynthesis occurs. As a practical matter, however, the concentration available to terrestrial plants is simply that found in the atmosphere: 0.035%.</p> <span id="Van_Niel"></span><span id="Van_Niel"></span><h2 class="lt-bio-4635">Van Niel</h2> <p class="lt-bio-4635">It was the American microbiologist Van Niel who first glimpsed the role that light plays in photosynthesis. He studied photosynthesis in purple sulfur bacteria. These microorganisms synthesize glucose from CO<sub><small><font size="1">2</font></small></sub> as do green plants, and they need light to do so. Water, however, is not the starting material. Instead they use hydrogen sulfide (H<sub><small><font size="1">2</font></small></sub>S). Furthermore, no oxygen is liberated during this photosynthesis but rather elemental sulfur. Van Niel reasoned that the action of light caused a decomposition of H<sub><small><font size="1">2</font></small></sub>S into hydrogen and sulfur atoms. 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href="#MJX-100-NCM-N-29"></use></g></g></g></g></svg></g></g></g></g></svg><mjx-assistive-mml unselectable="on" display="block"><math data-latex="\ce{6CO2 + 12H2O → C6H12O6 + 6 H2O + 6O2}" display="block" data-semantic-structure="(74 (73 (69 (66 (63 (60 5 59 (10 7 9)) 11 (62 12 61 (58 14 57 (56 (17 15 16) 55 18)))) 19 (65 (23 20 22) 64 (54 24 53 (52 (27 25 26) 51 (31 28 30))))) 32 (68 33 67 (50 35 49 (48 (38 36 37) 47 39))) 40 (71 41 70 (46 43 45))) (4 (3 0 1 2))))"><mtable displaystyle="true" 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}}}" data-semantic-type="multiline" data-semantic-role="unknown" data-semantic-annotation="depth:1" data-semantic-="" data-semantic-children="73" data-semantic-attributes="latex:\ce{6CO2 + 12H2O → C6H12O6 + 6 H2O + 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{4.9.3}" 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{4.9.3}" data-semantic-level-number="4" data-speech-node="true">4.9.3</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" data-semantic-annotation="depth:6" data-semantic-="" data-semantic-parent="65" data-semantic-added="true" data-semantic-operator="infixop,⁢" data-semantic-level-number="5" data-speech-node="true">⁢</mo><mrow data-semantic-added="true" data-semantic-type="appl" data-semantic-role="simple function" data-semantic-annotation="depth:6" data-semantic-="" data-semantic-children="24,52" data-semantic-content="53,24" data-semantic-parent="65" data-semantic-owns="24 53 52" data-semantic-level-number="5" 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:7" data-semantic-="" data-semantic-parent="54" data-semantic-attributes="latex:\mathrm{H};texclass:ORD" data-semantic-operator="appl" data-semantic-level-number="6" data-speech-node="true">H</mi></mrow><mo data-semantic-type="punctuation" data-semantic-role="application" data-semantic-annotation="nemeth:number;depth:7" data-semantic-="" data-semantic-parent="54" data-semantic-added="true" data-semantic-operator="appl" data-semantic-level-number="6" data-speech-node="true">⁡</mo><mrow data-semantic-added="true" data-semantic-type="infixop" data-semantic-role="implicit" data-semantic-annotation="clearspeak:unit;depth:7" data-semantic-="" data-semantic-children="27,31" data-semantic-content="51" data-semantic-parent="54" data-semantic-owns="27 51 31" data-semantic-level-number="6" data-speech-node="true"><msub data-latex="{\vphantom{A}}_{\smash[t]{12}}" data-semantic-type="subscript" data-semantic-role="unknown" data-semantic-annotation="depth:8" data-semantic-="" data-semantic-children="25,26" data-semantic-parent="52" data-semantic-attributes="latex:{\vphantom{A}}_{\smash[t]{12}}" data-semantic-owns="25 26" data-semantic-level-number="7" 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: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" data-speech-node="true">O</mi></mrow></mrow></mrow></mrow></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:9" data-semantic-="" data-semantic-parent="31" data-semantic-attributes="latex:{\smash[t]{6}};texclass:ORD" data-semantic-level-number="8" data-speech-node="true">6</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:4" data-semantic-="" data-semantic-parent="69" data-semantic-attributes="latex:+" data-semantic-operator="infixop,+" data-semantic-level-number="3" 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:4" data-semantic-="" data-semantic-children="33,50" data-semantic-content="67" data-semantic-parent="69" data-semantic-owns="33 67 50" data-semantic-level-number="3" 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;depth:5" data-semantic-="" data-semantic-parent="68" data-semantic-attributes="latex:6" data-semantic-level-number="4" data-speech-node="true">6</mn><mspace width="0.167em" data-latex="\,"></mspace><mo data-semantic-type="operator" data-semantic-role="multiplication" data-semantic-annotation="depth:5" data-semantic-="" data-semantic-parent="68" data-semantic-added="true" data-semantic-operator="infixop,⁢" data-semantic-level-number="4" data-speech-node="true">⁢</mo><mrow data-semantic-added="true" data-semantic-type="appl" data-semantic-role="simple function" data-semantic-annotation="depth:5" data-semantic-="" data-semantic-children="35,48" data-semantic-content="49,35" data-semantic-parent="68" data-semantic-owns="35 49 48" data-semantic-level-number="4" 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:6" data-semantic-="" data-semantic-parent="50" data-semantic-attributes="latex:\mathrm{H};texclass:ORD" data-semantic-operator="appl" data-semantic-level-number="5" data-speech-node="true">H</mi></mrow><mo data-semantic-type="punctuation" data-semantic-role="application" data-semantic-annotation="nemeth:number;depth:6" data-semantic-="" data-semantic-parent="50" data-semantic-added="true" data-semantic-operator="appl" data-semantic-level-number="5" data-speech-node="true">⁡</mo><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="38,39" data-semantic-content="47" data-semantic-parent="50" data-semantic-owns="38 47 39" data-semantic-level-number="5" data-speech-node="true"><msub data-latex="{\vphantom{A}}_{\smash[t]{2}}" data-semantic-type="subscript" data-semantic-role="unknown" data-semantic-annotation="depth:7" data-semantic-="" data-semantic-children="36,37" data-semantic-parent="48" data-semantic-attributes="latex:{\vphantom{A}}_{\smash[t]{2}}" data-semantic-owns="36 37" data-semantic-level-number="6" 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:8" data-semantic-="" data-semantic-parent="38" data-semantic-attributes="latex:{\vphantom{A}};texclass:ORD" data-semantic-level-number="7" 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:8" data-semantic-="" data-semantic-parent="38" data-semantic-attributes="latex:{\smash[t]{2}};texclass:ORD" data-semantic-level-number="7" data-speech-node="true">2</mn></mpadded></mrow></mrow></msub><mo data-semantic-type="operator" data-semantic-role="multiplication" data-semantic-annotation="depth:7" data-semantic-="" data-semantic-parent="48" data-semantic-added="true" data-semantic-operator="infixop,⁢" data-semantic-level-number="6" 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:7" data-semantic-="" data-semantic-parent="48" data-semantic-attributes="latex:\mathrm{O};texclass:ORD" data-semantic-level-number="6" data-speech-node="true">O</mi></mrow></mrow></mrow></mrow><mrow data-mjx-texclass="ORD" data-latex="{}"></mrow><mo data-latex="+" data-semantic-type="operator" data-semantic-role="addition" data-semantic-annotation="depth:4" data-semantic-="" data-semantic-parent="69" data-semantic-attributes="latex:+" data-semantic-operator="infixop,+" data-semantic-level-number="3" 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:4" data-semantic-="" data-semantic-children="41,46" data-semantic-content="70" data-semantic-parent="69" data-semantic-owns="41 70 46" data-semantic-level-number="3" 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;depth:5" data-semantic-="" data-semantic-parent="71" data-semantic-attributes="latex:6" data-semantic-level-number="4" data-speech-node="true">6</mn><mspace width="0.167em" data-latex="\," data-semantic-type="operator" data-semantic-role="space" data-semantic-annotation="depth:5" data-semantic-="" data-semantic-parent="71" data-semantic-operator="infixop,⁢" data-semantic-level-number="4" data-speech-node="true"></mspace><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: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 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:6" data-semantic-="" data-semantic-parent="46" data-semantic-attributes="latex:{\smash[t]{2}};texclass:ORD" data-semantic-level-number="5" data-speech-node="true">2</mn></mpadded></mrow></mrow></msub></mrow></mtd></mlabeledtr></mtable></math></mjx-assistive-mml></mjx-container></p> <p class="lt-bio-4635">In science, a theory should be testable. By deduction, one can make a prediction of how a particular experiment will come out if the theory is sound. In this case, the crucial experiments needed to test the two theories had to await the time when the growth of atomic research made it possible to produce isotopes other than those found naturally or in greater concentrations than are found naturally.</p> <span id="Samuel_Ruben"></span><span id="Samuel_Ruben"></span><h2 class="lt-bio-4635">Samuel Ruben</h2> <p class="lt-bio-4635">In air, water and other natural materials containing oxygen, 99.76% of the oxygen atoms are <sup><small><font size="1">16</font></small></sup>O and only 0.20% of them are the heavier isotope <sup><small><font size="1">18</font></small></sup>O. In 1941, Samuel Ruben and his coworkers at the University of California were able to prepare specially "labeled" water in which the 0.85% of the molecules contained <sup><small><font size="1">18</font></small></sup>O atoms. When this water was supplied to a suspension of photosynthesizing algae, the proportion of <sup><small><font size="1">18</font></small></sup>O in the oxygen gas that was evolved was 0.85%, the same as that of the water supplied, and not simply the 0.20% found in all natural samples of oxygen (and its compounds like CO<sub><small><font size="1">2</font></small></sub>).</p> <table class="mt-responsive-table"> <tbody> <tr> <td class="mt-align-center mt-noheading lt-bio-4635" colspan="2"> </td> <td class="mt-align-center mt-noheading lt-bio-4635" colspan="3">% <sup><small><font size="1">18</font></small></sup>O FOUND IN</td> </tr> <tr> <td class="mt-align-center mt-noheading lt-bio-4635">EXPERIMENT</td> <td class="mt-align-center mt-noheading lt-bio-4635"> </td> <td class="mt-align-center mt-noheading lt-bio-4635">H<sub><small><font size="1">2</font></small></sub>O</td> <td class="mt-align-center mt-noheading lt-bio-4635">CO<sub><small><font size="1">2</font></small></sub></td> <td class="mt-align-center mt-noheading lt-bio-4635">O<sub><small><font size="1">2</font></small></sub></td> </tr> <tr> <td class="mt-align-center mt-noheading lt-bio-4635">1.</td> <td class="mt-align-center mt-noheading lt-bio-4635">START</td> <td class="mt-align-center mt-noheading lt-bio-4635">0.85</td> <td class="mt-align-center mt-noheading lt-bio-4635">0.20</td> <td class="mt-align-center mt-noheading lt-bio-4635">—</td> </tr> <tr> <td class="mt-align-center mt-noheading lt-bio-4635"> </td> <td class="mt-align-center mt-noheading lt-bio-4635">FINISH</td> <td class="mt-align-center mt-noheading lt-bio-4635">0.85</td> <td class="mt-align-center mt-noheading lt-bio-4635">0.61<span style="color:red;">*</span></td> <td class="mt-align-center mt-noheading lt-bio-4635">0.86</td> </tr> <tr> <td class="mt-align-center mt-noheading lt-bio-4635">2.</td> <td class="mt-align-center mt-noheading lt-bio-4635">START</td> <td class="mt-align-center mt-noheading lt-bio-4635">0.20</td> <td class="mt-align-center mt-noheading lt-bio-4635">0.68</td> <td class="mt-align-center mt-noheading lt-bio-4635">—</td> </tr> <tr> <td class="mt-align-center mt-noheading lt-bio-4635"> </td> <td class="mt-align-center mt-noheading lt-bio-4635">FINISH</td> <td class="mt-align-center mt-noheading lt-bio-4635">0.20</td> <td class="mt-align-center mt-noheading lt-bio-4635">0.57</td> <td class="mt-align-center mt-noheading lt-bio-4635">0.20</td> </tr> </tbody> </table> <p class="lt-bio-4635"><big><span style="color:red;"><font size="5">*</font></span></big> A non-biochemical exchange of oxygen atoms between the water and the bicarbonate ions used as a source of CO<sub><small><font size="1">2</font></small></sub> explains the uptake of the isotope by CO<sub><small><font size="1">2</font></small></sub> in the first experiment.</p> <p class="lt-bio-4635">These results clearly demonstrated that Senebier's interpretation was in error. If all the oxygen liberated during photosynthesis comes from the carbon dioxide, we would expect the oxygen evolved in Ruben's experiment to contain simply the 0.20% found naturally. If, on the other hand, both the carbon dioxide and the water contribute to the oxygen released, we would expect its isotopic composition to have been some intermediate figure. In fact, the isotopic composition of the evolved oxygen was the same as that of the water used.</p> <p class="lt-bio-4635">Ruben and his colleagues also prepared a source of carbon dioxide that was enriched in <sup><small><font size="1">18</font></small></sup>O atoms. When algae carried out photosynthesis using this material and <strong>natural</strong> water, the oxygen that was given off was not enriched in <sup><small><font size="1">18</font></small></sup>O. It contained simply the 0.20% <sup><small><font size="1">18</font></small></sup>O found in the natural water used. The heavy atoms presumably became incorporated in the other two products (carbohydrate and by-product water).</p> <p class="lt-bio-4635">These experiments lent great support to Van Niel's idea that one function of light in photosynthesis was the separation of the hydrogen and oxygen atoms of water molecules. 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