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src="//upload.wikimedia.org/wikipedia/commons/thumb/1/10/Marshall_Nirenberg_performing_experiment.jpg/250px-Marshall_Nirenberg_performing_experiment.jpg" decoding="async" width="250" height="312" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/10/Marshall_Nirenberg_performing_experiment.jpg/375px-Marshall_Nirenberg_performing_experiment.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/10/Marshall_Nirenberg_performing_experiment.jpg/500px-Marshall_Nirenberg_performing_experiment.jpg 2x" data-file-width="4728" data-file-height="5892" /></a><figcaption>Marshall Nirenberg</figcaption></figure><p>The <b>Nirenberg and Leder experiment</b> was a scientific experiment performed in 1964 by <a href="/wiki/Marshall_W._Nirenberg" class="mw-redirect" title="Marshall W. Nirenberg">Marshall W. Nirenberg</a> and <a href="/wiki/Philip_Leder" title="Philip Leder">Philip Leder</a>. The experiment elucidated the triplet nature of the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> and allowed the remaining ambiguous codons in the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> to be deciphered. </p><p>In this experiment, using a <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> binding <a href="/wiki/Assay" title="Assay">assay</a> called the <a href="#Experimental_work">triplet binding assay</a>, various combinations of <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a> were passed through a filter which contained ribosomes. Unique triplets promoted the binding of specific <a href="/wiki/TRNA" class="mw-redirect" title="TRNA">tRNAs</a> to the ribosome. By associating the tRNA with its specific <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a>, it was possible to determine the triplet mRNA sequence that coded for each amino acid. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Background">Background</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Nirenberg_and_Leder_experiment&action=edit&section=1" title="Edit section: Background"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Oswald_Avery" title="Oswald Avery">Oswald Avery</a> discovered that the substance responsible for producing inheritable change in the disease-causing bacteria was neither a protein nor a lipid, rather deoxyribonucleic acid (<a href="/wiki/DNA" title="DNA">DNA</a>). He and his colleagues <a href="/wiki/Colin_Munro_MacLeod" title="Colin Munro MacLeod">Colin MacLeod</a> and <a href="/wiki/Maclyn_McCarty" title="Maclyn McCarty">Maclyn McCarty</a> suggested that DNA was responsible for transferring genetic information. Later, <a href="/wiki/Erwin_Chargaff" title="Erwin Chargaff">Erwin Chargaff</a> discovered that the makeup of DNA differs from one species to another. These experiments helped pave the way for the discovery of the structure of DNA. In 1953, with the help of <a href="/wiki/Maurice_Wilkins" title="Maurice Wilkins">Maurice Wilkins</a> and <a href="/wiki/Rosalind_Franklin" title="Rosalind Franklin">Rosalind Franklin</a>’s <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a>, <a href="/wiki/James_D._Watson" class="mw-redirect" title="James D. Watson">James Watson</a> and <a href="/wiki/Francis_Crick" title="Francis Crick">Francis Crick</a> proposed DNA is structured as a <a href="/wiki/Double_helix" class="mw-redirect" title="Double helix">double helix</a>.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>In the 1960s, one main DNA mystery scientists needed to figure out was in <a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">translation</a> how many bases would be in each code word, or <a href="/wiki/Codon" class="mw-redirect" title="Codon">codon</a>. Scientists knew there were a total of four bases (<a href="/wiki/Guanine" title="Guanine">guanine</a>, <a href="/wiki/Cytosine" title="Cytosine">cytosine</a>, <a href="/wiki/Adenine" title="Adenine">adenine</a>, and <a href="/wiki/Thymine" title="Thymine">thymine</a>). They also knew that were 20 known <a href="/wiki/Amino_acids" class="mw-redirect" title="Amino acids">amino acids</a>. <a href="/wiki/George_Gamow" title="George Gamow">George Gamow</a> suggested that the genetic code was made of three nucleotides per amino acid. He reasoned that because there are 20 amino acids and only four bases, the coding units could not be single (4 combinations) or pairs (only 16 combinations). Rather, he thought triplets (64 possible combinations) were the coding unit of the genetic code. However, he proposed that the triplets were overlapping and <a href="/wiki/Genetic_code#Degeneracy" title="Genetic code">non-degenerate</a>.<sup id="cite_ref-Leavitt_craze_2-0" class="reference"><a href="#cite_note-Leavitt_craze-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Seymour_Benzer" title="Seymour Benzer">Seymour Benzer</a> in the late 1950s had developed an assay using phage mutations which provided the first detailed linearly structured map of a genetic region. Crick felt he could use mutagenesis and genetic recombination phage to further delineate the nature of the genetic code.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> In the <a href="/wiki/Crick,_Brenner_et_al._experiment" title="Crick, Brenner et al. experiment">Crick, Brenner et al. experiment</a>, using these phages, the triplet nature of the genetic code was confirmed. They used <a href="/wiki/Frameshift_mutation" title="Frameshift mutation">frameshift mutations</a> and a process called <a href="/wiki/Mutation#Classification_of_mutation_types" title="Mutation">reversions</a>, to add and delete various numbers of nucleotides.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> When a nucleotide triplet was added to or deleted from the DNA sequence, the encoded protein was minimally affected. Thus, they concluded that the genetic code is a triplet code because it did not cause a frameshift in the reading frame.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> They correctly concluded that the code is degenerate, that triplets are not overlapping, and that each nucleotide sequence is read from a specific starting point. </p> <div class="mw-heading mw-heading2"><h2 id="Experimental_work">Experimental work</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Nirenberg_and_Leder_experiment&action=edit&section=2" title="Edit section: Experimental work"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:06_multi_pu.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7e/06_multi_pu.jpg/175px-06_multi_pu.jpg" decoding="async" width="175" height="114" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7e/06_multi_pu.jpg/263px-06_multi_pu.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7e/06_multi_pu.jpg/350px-06_multi_pu.jpg 2x" data-file-width="400" data-file-height="261" /></a><figcaption>The Multi-plater, developed by Leder, helped speed up the process of deciphering the genetic code.<sup id="cite_ref-Leavitt_Instruments_6-0" class="reference"><a href="#cite_note-Leavitt_Instruments-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The very first amino acid codon (UUU encoding phenylalanine) was deciphered by Nirenberg and his postdoc <a href="/wiki/Heinrich_Matthaei" class="mw-redirect" title="Heinrich Matthaei">Heinrich Matthaei</a> (see <i><a href="/wiki/Nirenberg_and_Matthaei_experiment" title="Nirenberg and Matthaei experiment">Nirenberg and Matthaei experiment</a></i>) using long synthetic RNA. However, when similar RNAs are made containing more than one RNA base, the order of the bases was random. For example, a long RNA could be made that had a ratio of C to U of 2:1, and so would contain codons CCU, CUC, UCC at high frequency. When translated by ribosomes, this would produce a protein containing the amino acids proline, leucine, and serine; but it was not possible to say which codon matched which amino acid.<sup id="cite_ref-Judson_H._1996_7-0" class="reference"><a href="#cite_note-Judson_H._1996-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>Instead, Nirenberg's group turned to very short synthetic RNAs. They found that the trinucleotide UUU (which is the codon for phenylalanine), was able to cause specific association of phenylalanine-charged tRNA with ribosomes. This association could be detected by passing the mixture through a nitrocellulose filter: the filter captures ribosomes but not free tRNA; however if tRNA was associated with the ribosome, it would also be captured (along with the radioactive phenylalanine attached to the tRNA). They similarly found that trinucleotides AAA or CCC caused ribosome association of lysine-tRNA or proline-tRNA, respectively. <sup id="cite_ref-Nirenberg_1964_8-0" class="reference"><a href="#cite_note-Nirenberg_1964-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>So an experimental plan was clear: synthesize all 64 different trinucleotide combinations, and use the filter assay with tRNAs charged with all 20 amino acids, to see which amino acid associated with which trinucleotide. However, obtaining pure trinucleotides with mixed base sequences, for example GUU, was a daunting challenge. Leder's pioneering studies used trinucleotides made by breaking down long random poly-GU RNA with nuclease and purifying specific trinucleotides by <a href="/wiki/Paper_chromatography" title="Paper chromatography">paper chromatography</a>:<sup id="cite_ref-Nirenberg_1964_8-1" class="reference"><a href="#cite_note-Nirenberg_1964-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> he determined that GUU, UGU, and UUG encoded the amino acids valine,<sup id="cite_ref-Leder_1964a_9-0" class="reference"><a href="#cite_note-Leder_1964a-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> cysteine and leucine,<sup id="cite_ref-Leder_1964b_10-0" class="reference"><a href="#cite_note-Leder_1964b-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> respectively. Subsequently, Nirenberg's group constructed trinucleotides by using DNA polymerases coupled with nucleotides and RNA polymerases to create the long random poly-GU RNA as well as artificially replicate the purified trinucleotides. Once high enough concentrations of mRNA were produced, degradation and reformation of polymerase products was accomplished through enzymatic processes. For example, AGU could be made from AG and U with <a href="/wiki/Polynucleotide_phosphorylase" title="Polynucleotide phosphorylase">polynucleotide phosphorylase</a>; UAG could be made from AG and U with <a href="/wiki/Ribonuclease_A" class="mw-redirect" title="Ribonuclease A">ribonuclease A</a> in a high concentration of methanol.<sup id="cite_ref-Nirenberg_2004_11-0" class="reference"><a href="#cite_note-Nirenberg_2004-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Nirenberg's postdoc <a href="/wiki/Merton_Bernfield" title="Merton Bernfield">Merton Bernfield</a> used these techniques to determine that UUU and UUC encode phenylalanine, UCU and UCC encode serine, and CCC and CCU encode proline, highlighting a pattern in the way the genetic code redundantly encodes amino acids.<sup id="cite_ref-Bernfield_1964_12-0" class="reference"><a href="#cite_note-Bernfield_1964-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Many others in the Nirenberg lab and at NIH contributed to the full decipherment of the genetic code.<sup id="cite_ref-Nirenberg_2004_11-1" class="reference"><a href="#cite_note-Nirenberg_2004-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Reception_and_legacy">Reception and legacy</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Nirenberg_and_Leder_experiment&action=edit&section=3" title="Edit section: Reception and legacy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:06_chart_pu3.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/21/06_chart_pu3.png/250px-06_chart_pu3.png" decoding="async" width="250" height="219" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/21/06_chart_pu3.png/375px-06_chart_pu3.png 1.5x, //upload.wikimedia.org/wikipedia/commons/2/21/06_chart_pu3.png 2x" data-file-width="400" data-file-height="350" /></a><figcaption> Genetic Code Chart</figcaption></figure><p>By the Cold Spring Harbor Symposium of 1966, between Nirenberg and Khorana the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a> was almost completely decoded. Nirenberg was awarded the 1968 <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a>. He shared the award with <a href="/wiki/Har_Gobind_Khorana" title="Har Gobind Khorana">Har Gobind Khorana</a> of the <a href="/wiki/University_of_Wisconsin" class="mw-redirect" title="University of Wisconsin">University of Wisconsin</a> and Robert W. Holley of the Salk Institute. Working independently, Khorana had mastered the synthesis of nucleic acids, and Holley had discovered the exact chemical structure of transfer-RNA. <br /> </p><p><i>The New York Times</i> said of Nirenberg's work that "the science of biology has reached a new frontier," leading to "a revolution far greater in its potential significance than the atomic or hydrogen bomb." Most of the scientific community saw these experiments as highly important and beneficial. However, there were some who were concerned with the new era of <a href="/wiki/Molecular_genetics" title="Molecular genetics">molecular genetics</a>. For example, <a href="/wiki/Arne_Tiselius" title="Arne Tiselius">Arne Tiselius</a>, the 1948 Nobel Laureate in Chemistry, asserted that knowledge of the genetic code could "lead to methods of tampering with life, of creating new diseases, of controlling minds, of influencing heredity, even perhaps in certain desired directions."<sup id="cite_ref-Fee_public_13-0" class="reference"><a href="#cite_note-Fee_public-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Nirenberg_and_Leder_experiment&action=edit&section=4" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFRussell_P.2010" class="citation book cs1">Russell P. 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The Effect of Trinucleotides upon the Binding of sRNA to Ribosomes". <i>Science</i>. <b>145</b> (3639): 1399–1407. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.145.3639.1399">10.1126/science.145.3639.1399</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/14172630">14172630</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:7127820">7127820</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=RNA+Codewords+and+Protein+Synthesis.+The+Effect+of+Trinucleotides+upon+the+Binding+of+sRNA+to+Ribosomes&rft.volume=145&rft.issue=3639&rft.pages=1399-1407&rft.date=1964&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A7127820%23id-name%3DS2CID&rft_id=info%3Apmid%2F14172630&rft_id=info%3Adoi%2F10.1126%2Fscience.145.3639.1399&rft.au=Philip+Leder&rft.au=Marshall+Nirenberg&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANirenberg+and+Leder+experiment" class="Z3988"></span></span> </li> <li id="cite_note-Leder_1964a-9"><span class="mw-cite-backlink"><b><a href="#cite_ref-Leder_1964a_9-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLeder_P.Nirenberg_M.W.1964" class="citation journal cs1">Leder P. & Nirenberg M.W. (1964). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC300293">"RNA Codewords and Protein Synthesis, II. Nucleotide Sequence of a Valine RNA Codeword"</a>. <i>PNAS</i>. <b>52</b> (2): 420–427. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1964PNAS...52..420L">1964PNAS...52..420L</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.52.2.420">10.1073/pnas.52.2.420</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC300293">300293</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/14206609">14206609</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PNAS&rft.atitle=RNA+Codewords+and+Protein+Synthesis%2C+II.+Nucleotide+Sequence+of+a+Valine+RNA+Codeword&rft.volume=52&rft.issue=2&rft.pages=420-427&rft.date=1964&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC300293%23id-name%3DPMC&rft_id=info%3Apmid%2F14206609&rft_id=info%3Adoi%2F10.1073%2Fpnas.52.2.420&rft_id=info%3Abibcode%2F1964PNAS...52..420L&rft.au=Leder+P.&rft.au=Nirenberg+M.W.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC300293&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANirenberg+and+Leder+experiment" class="Z3988"></span></span> </li> <li id="cite_note-Leder_1964b-10"><span class="mw-cite-backlink"><b><a href="#cite_ref-Leder_1964b_10-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLeder_P.Nirenberg_M.W.1964" class="citation journal cs1">Leder P. & Nirenberg M.W. (1964). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC300480">"RNA Codewords and Protein Synthesis, III. On the Nucleotide Sequence of a Cysteine and a Leucine RNA Codeword"</a>. <i>PNAS</i>. <b>52</b> (6): 1521–1529. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1964PNAS...52.1521L">1964PNAS...52.1521L</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.52.6.1521">10.1073/pnas.52.6.1521</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC300480">300480</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/14243527">14243527</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PNAS&rft.atitle=RNA+Codewords+and+Protein+Synthesis%2C+III.+On+the+Nucleotide+Sequence+of+a+Cysteine+and+a+Leucine+RNA+Codeword&rft.volume=52&rft.issue=6&rft.pages=1521-1529&rft.date=1964&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC300480%23id-name%3DPMC&rft_id=info%3Apmid%2F14243527&rft_id=info%3Adoi%2F10.1073%2Fpnas.52.6.1521&rft_id=info%3Abibcode%2F1964PNAS...52.1521L&rft.au=Leder+P.&rft.au=Nirenberg+M.W.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC300480&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANirenberg+and+Leder+experiment" class="Z3988"></span></span> </li> <li id="cite_note-Nirenberg_2004-11"><span class="mw-cite-backlink">^ <a href="#cite_ref-Nirenberg_2004_11-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Nirenberg_2004_11-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNirenberg,_Marshall2004" class="citation journal cs1">Nirenberg, Marshall (2004). "Historical review: Deciphering the genetic code – a personal account". <i>Trends in Biochemical Sciences</i>. <b>29</b> (1): 46–54. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.tibs.2003.11.009">10.1016/j.tibs.2003.11.009</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/14729332">14729332</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Trends+in+Biochemical+Sciences&rft.atitle=Historical+review%3A+Deciphering+the+genetic+code+%E2%80%93+a+personal+account&rft.volume=29&rft.issue=1&rft.pages=46-54&rft.date=2004&rft_id=info%3Adoi%2F10.1016%2Fj.tibs.2003.11.009&rft_id=info%3Apmid%2F14729332&rft.au=Nirenberg%2C+Marshall&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANirenberg+and+Leder+experiment" class="Z3988"></span></span> </li> <li id="cite_note-Bernfield_1964-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-Bernfield_1964_12-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBernfield_M.R.Nirenberg_M.W.1965" class="citation journal cs1">Bernfield M.R. & Nirenberg M.W. (1965). "RNA Codewords and Protein Synthesis. The Nucleotide Sequences of Multiple Codewords for Phenylalanine, Serine, Leucine, and Proline". <i>Science</i>. <b>147</b> (3657): 479–484. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.147.3657.479">10.1126/science.147.3657.479</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/14237203">14237203</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=RNA+Codewords+and+Protein+Synthesis.+The+Nucleotide+Sequences+of+Multiple+Codewords+for+Phenylalanine%2C+Serine%2C+Leucine%2C+and+Proline&rft.volume=147&rft.issue=3657&rft.pages=479-484&rft.date=1965&rft_id=info%3Adoi%2F10.1126%2Fscience.147.3657.479&rft_id=info%3Apmid%2F14237203&rft.au=Bernfield+M.R.&rft.au=Nirenberg+M.W.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANirenberg+and+Leder+experiment" class="Z3988"></span></span> </li> <li id="cite_note-Fee_public-13"><span class="mw-cite-backlink"><b><a href="#cite_ref-Fee_public_13-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFee2000" class="citation web cs1">Fee, E. (2000). <a rel="nofollow" class="external text" href="https://profiles.nlm.nih.gov/spotlight/jj/feature/publicreaction">"Profiles in Science: The Marshall W. Nirenberg Papers. Public Reaction"</a>. National Library of Medicine. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20200409213318/https://profiles.nlm.nih.gov/spotlight/jj/feature/publicreaction">Archived</a> from the original on 9 April 2020<span class="reference-accessdate">. Retrieved <span class="nowrap">9 April</span> 2020</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Profiles+in+Science%3A+The+Marshall+W.+Nirenberg+Papers.+Public+Reaction&rft.pub=National+Library+of+Medicine&rft.date=2000&rft.aulast=Fee&rft.aufirst=E.&rft_id=https%3A%2F%2Fprofiles.nlm.nih.gov%2Fspotlight%2Fjj%2Ffeature%2Fpublicreaction&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANirenberg+and+Leder+experiment" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Nirenberg_and_Leder_experiment&action=edit&section=5" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://id.lib.harvard.edu/aleph/011455194/catalog">Philip Leder's laboratory notebooks, December 1963 to August 1965</a>, can be found at The Center for the History of Medicine at the Countway Library, Harvard Medical School. Digital surrogates are also available through Harvard's HOLLIS catalog.</li></ul> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Nirenberg_and_Leder_experiment&action=edit&section=6" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Crick,_Brenner_et_al._experiment" title="Crick, Brenner et al. experiment">Crick, Brenner et al. experiment</a></li> <li><a href="/wiki/Nirenberg_and_Matthaei_experiment" title="Nirenberg and Matthaei experiment">Nirenberg and Matthaei experiment</a></li> <li><a href="/wiki/List_of_experiments" title="List of experiments">List of experiments</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output 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href="/wiki/History_of_agricultural_science" title="History of agricultural science">Agricultural science</a></li> <li><a href="/wiki/History_of_anatomy" title="History of anatomy">Anatomy</a></li> <li><a href="/wiki/History_of_biochemistry" title="History of biochemistry">Biochemistry</a></li> <li><a href="/wiki/History_of_biotechnology" title="History of biotechnology">Biotechnology</a></li> <li><a href="/wiki/History_of_botany" title="History of botany">Botany</a></li> <li><a href="/wiki/History_of_ecology" title="History of ecology">Ecology</a></li> <li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Evolutionary thought</a></li> <li><a href="/wiki/History_of_genetics" title="History of genetics">Genetics</a></li> <li><a href="/wiki/History_of_geology" title="History of geology">Geology</a></li> <li><a href="/wiki/Timeline_of_immunology" title="Timeline of immunology">Immunology</a></li> <li><a href="/wiki/History_of_medicine" title="History 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