Revolutionizing Medicine with Monoclonal Antibodies | Scientists and Research

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The module describes Milstein’s life as a student in Argentina during the regime of Juan Peron and his later research as he became a world-class scientist, winning the Nobel Prize in 1984."> <meta name="keywords" content="César Milstein, Milstein, hybridoma, antibodies, antigen, mutation"> <meta name="viewport" content="width=device-width, initial-scale=1.0, shrink-to-fit=no"> <meta name="msvalidate.01" content="D8E20F39AD48052260032E56DE409970"> <script type="application/ld+json"> { "@context": "http://schema.org/", "@type": "Article", "mainEntityOfPage": { "@type": "WebPage", "@id": "https://visionlearning.com/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220" }, "name": "Revolutionizing Medicine with Monoclonal Antibodies", "headline": "Revolutionizing Medicine with Monoclonal Antibodies: The work of César Milstein", "author": { "@type": "Person", "name": "David Warmflash, MD" }, "datePublished": "2015-07-05 09:29:13", "dateModified": "2017-02-12T08:30:00+05:00", "image": { "@type": "ImageObject", "url": "/img/library/moduleImages/featured_image_220-23061210064208.jpeg", "width": 696, "height": 464 }, "publisher": { "@type": "Organization", "name": "Visionlearning, Inc.", "logo": { "@type": "ImageObject", "url": "http://visionlearning.com/images/logo.png", "width": 278, "height": 60 } }, "description": "From home pregnancy tests to treatment for Ebola, the discovery of monoclonal antibodies has greatly advanced science and medicine. This module traces the work of immunologist César Milstein, who successfully created “hybridoma” cells capable of producing specific antibodies in mass quantities. The module describes Milstein’s life as a student in Argentina during the regime of Juan Peron and his later research as he became a world-class scientist, winning the Nobel Prize in 1984.", "keywords": "César Milstein, Milstein, hybridoma, antibodies, antigen, mutation", "inLanguage": { "@type": "Language", "name": "English", "alternateName": "en" }, "copyrightHolder": { "@type": "Organization", "name": "Visionlearning, Inc." }, "copyrightYear": "2015"} </script> <meta property="og:url" content="https://visionlearning.com/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220"> <meta property="og:title" content="Revolutionizing Medicine with Monoclonal Antibodies | Scientists and Research | Visionlearning" /> <meta property="og:type" content="website"> <meta property="og:site_name" content="Visionlearning"> <meta property="og:description" content="From home pregnancy tests to treatment for Ebola, the discovery of monoclonal antibodies has greatly advanced science and medicine. This module traces the work of immunologist César Milstein, who successfully created “hybridoma” cells capable of producing specific antibodies in mass quantities. 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<li><a href="/en/library/environmental-science/61/ecosystem-services/279">Ecosystem Services</a></li> <li><a href="/en/library/environmental-science/61/population-biology/287">Population Biology</a></li> </ul> </div> <button class="accordion__button" id="acc-button-earth-cycles" data-accordion="button" aria-controls="acc-panel-earth-cycles" aria-expanded="false"> <span class="accordion__button__label"> Earth Cycles </span> </button> <div class="accordion__panel" id="acc-panel-earth-cycles" data-accordion="panel" aria-labelledby="acc-button-earth-cycles" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/environmental-science/61/the-nitrogen-cycle/98">The Nitrogen Cycle</a></li> <li><a href="/en/library/environmental-science/61/the-carbon-cycle/95">The Carbon Cycle</a></li> <li><a href="/en/library/environmental-science/61/the-phosphorus-cycle/197">The Phosphorus Cycle</a></li> </ul> </div> <button class="accordion__button" id="acc-button-scientific-research" data-accordion="button" aria-controls="acc-panel-scientific-research" aria-expanded="false"> <span class="accordion__button__label"> Scientific Research </span> </button> <div class="accordion__panel" id="acc-panel-scientific-research" data-accordion="panel" aria-labelledby="acc-button-scientific-research" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/environmental-science/61/collaborative-research-in-the-arctic-towards-understanding-climate-change/183">Collaborative Research in the Arctic Towards Understanding Climate Change</a></li> <li><a href="/en/library/environmental-science/61/atmospheric-chemistry-research-that-changed-global-policy/211">Atmospheric Chemistry Research that Changed Global Policy</a></li> </ul> </div> </div> </div> <button class="accordion__button" id="acc-button-general-science" data-accordion="button" aria-controls="acc-panel-general-science" aria-expanded="false"> <span class="accordion__button__label"> General Science </span> 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aria-labelledby="acc-button-measurement" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/general-science/3/the-metric-system/47">The Metric System</a></li> </ul> </div> <button class="accordion__button" id="acc-button-physical-properties" data-accordion="button" aria-controls="acc-panel-physical-properties" aria-expanded="false"> <span class="accordion__button__label"> Physical Properties </span> </button> <div class="accordion__panel" id="acc-panel-physical-properties" data-accordion="panel" aria-labelledby="acc-button-physical-properties" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/general-science/3/temperature/48">Temperature</a></li> <li><a href="/en/library/general-science/3/density-and-buoyancy/37">Density and Buoyancy</a></li> </ul> </div> </div> </div> <button class="accordion__button" id="acc-button-math-in-science" data-accordion="button" aria-controls="acc-panel-math-in-science" aria-expanded="false"> <span 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href="/en/library/math-in-science/62/exponential-equations-ii/210">Exponential Equations II</a></li> <li><a href="/en/library/math-in-science/62/scientific-notation/250">Scientific Notation</a></li> <li><a href="/en/library/math-in-science/62/measurement/257">Measurement</a></li> </ul> </div> <button class="accordion__button" id="acc-button-statistics" data-accordion="button" aria-controls="acc-panel-statistics" aria-expanded="false"> <span class="accordion__button__label"> Statistics </span> </button> <div class="accordion__panel" id="acc-panel-statistics" data-accordion="panel" aria-labelledby="acc-button-statistics" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/math-in-science/62/introduction-to-descriptive-statistics/218">Introduction to Descriptive Statistics</a></li> <li><a href="/en/library/math-in-science/62/introduction-to-inferential-statistics/224">Introduction to Inferential Statistics</a></li> <li><a href="/en/library/math-in-science/62/statistical-techniques/239">Statistical Techniques</a></li> </ul> </div> <button class="accordion__button" id="acc-button-trigonometric-functions" data-accordion="button" aria-controls="acc-panel-trigonometric-functions" aria-expanded="false"> <span class="accordion__button__label"> Trigonometric Functions </span> </button> <div class="accordion__panel" id="acc-panel-trigonometric-functions" data-accordion="panel" aria-labelledby="acc-button-trigonometric-functions" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/math-in-science/62/wave-mathematics/131">Wave Mathematics</a></li> </ul> </div> </div> </div> <button class="accordion__button" id="acc-button-physics" data-accordion="button" aria-controls="acc-panel-physics" aria-expanded="false"> <span class="accordion__button__label"> Physics </span> </button> <div class="accordion__panel" id="acc-panel-physics" data-accordion="panel" aria-labelledby="acc-button-physics" role="region"> <div class="accordion accordion--secondary"> <button class="accordion__button" id="acc-button-light-and-optics" data-accordion="button" aria-controls="acc-panel-light-and-optics" aria-expanded="false"> <span class="accordion__button__label"> Light and Optics </span> </button> <div class="accordion__panel" id="acc-panel-light-and-optics" data-accordion="panel" aria-labelledby="acc-button-light-and-optics" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/physics/24/the-nature-of-light/132">The Nature of Light</a></li> <li><a href="/en/library/physics/24/electromagnetism-and-light/138">Electromagnetism and Light</a></li> </ul> </div> <button class="accordion__button" id="acc-button-mechanics" data-accordion="button" aria-controls="acc-panel-mechanics" aria-expanded="false"> <span class="accordion__button__label"> Mechanics </span> </button> <div class="accordion__panel" id="acc-panel-mechanics" data-accordion="panel" aria-labelledby="acc-button-mechanics" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/physics/24/defining-energy/199">Defining Energy</a></li> <li><a href="/en/library/physics/24/waves-and-wave-motion/102">Waves and Wave Motion</a></li> <li><a href="/en/library/physics/24/gravity/118">Gravity</a></li> <li><a href="/en/library/physics/24/thermodynamics-i/200">Thermodynamics I</a></li> </ul> </div> </div> </div> <button class="accordion__button" id="acc-button-process-of-science" data-accordion="button" aria-controls="acc-panel-process-of-science" aria-expanded="false"> <span class="accordion__button__label"> Process of Science </span> </button> <div class="accordion__panel" id="acc-panel-process-of-science" data-accordion="panel" aria-labelledby="acc-button-process-of-science" role="region"> <div class="accordion accordion--secondary"> <button class="accordion__button" id="acc-button-introduction" data-accordion="button" aria-controls="acc-panel-introduction" aria-expanded="false"> <span class="accordion__button__label"> Introduction </span> </button> <div class="accordion__panel" id="acc-panel-introduction" data-accordion="panel" aria-labelledby="acc-button-introduction" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/process-of-science/49/the-process-of-science/176">The Process of Science</a></li> </ul> </div> <button class="accordion__button" id="acc-button-the-culture-of-science" data-accordion="button" aria-controls="acc-panel-the-culture-of-science" aria-expanded="false"> <span class="accordion__button__label"> The Culture of Science </span> </button> <div class="accordion__panel" id="acc-panel-the-culture-of-science" data-accordion="panel" aria-labelledby="acc-button-the-culture-of-science" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/process-of-science/49/the-nature-of-scientific-knowledge/185">The Nature of Scientific Knowledge</a></li> <li><a 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href="/en/library/process-of-science/49/comparison-in-scientific-research/152">Comparison in Scientific Research</a></li> <li><a href="/en/library/process-of-science/49/modeling-in-scientific-research/153">Modeling in Scientific Research</a></li> </ul> </div> <button class="accordion__button" id="acc-button-data" data-accordion="button" aria-controls="acc-panel-data" aria-expanded="false"> <span class="accordion__button__label"> Data </span> </button> <div class="accordion__panel" id="acc-panel-data" data-accordion="panel" aria-labelledby="acc-button-data" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/process-of-science/49/data-analysis-and-interpretation/154">Data Analysis and Interpretation</a></li> <li><a href="/en/library/process-of-science/49/uncertainty-error-and-confidence/157">Uncertainty, Error, and Confidence</a></li> <li><a href="/en/library/process-of-science/49/statistics-in-science/155">Statistics in Science</a></li> <li><a href="/en/library/process-of-science/49/using-graphs-and-visual-data-in-science/156">Using Graphs and Visual Data in Science</a></li> </ul> </div> <button class="accordion__button" id="acc-button-scientific-communication" data-accordion="button" aria-controls="acc-panel-scientific-communication" aria-expanded="false"> <span class="accordion__button__label"> Scientific Communication </span> </button> <div class="accordion__panel" id="acc-panel-scientific-communication" data-accordion="panel" aria-labelledby="acc-button-scientific-communication" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/process-of-science/49/understanding-scientific-journals-and-articles/158">Understanding Scientific Journals and Articles</a></li> <li><a href="/en/library/process-of-science/49/utilizing-the-scientific-literature/173">Utilizing the Scientific Literature</a></li> <li><a href="/en/library/process-of-science/49/peer-review-in-scientific-publishing/159">Peer Review in Scientific Publishing</a></li> <li><a href="/en/library/process-of-science/49/the-how-and-why-of-scientific-meetings/186">The How and Why of Scientific Meetings</a></li> </ul> </div> </div> </div> <button class="accordion__button" id="acc-button-scientists-and-research" data-accordion="button" aria-controls="acc-panel-scientists-and-research" aria-expanded="false"> <span class="accordion__button__label"> Scientists and Research </span> </button> <div class="accordion__panel" id="acc-panel-scientists-and-research" data-accordion="panel" aria-labelledby="acc-button-scientists-and-research" role="region"> <div class="accordion accordion--secondary"> <button class="accordion__button" id="acc-button-scientific-research" data-accordion="button" aria-controls="acc-panel-scientific-research" aria-expanded="false"> <span class="accordion__button__label"> Scientific Research </span> </button> <div class="accordion__panel" id="acc-panel-scientific-research" data-accordion="panel" aria-labelledby="acc-button-scientific-research" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/scientists-and-research/58/collaborative-research-in-the-arctic-towards-understanding-climate-change/183">Collaborative Research in the Arctic Towards Understanding Climate Change</a></li> <li><a href="/en/library/scientists-and-research/58/from-stable-chromosomes-to-jumping-genes/184">From Stable Chromosomes to Jumping Genes</a></li> <li><a href="/en/library/scientists-and-research/58/an-elegant-experiment-to-test-the-process-of-dna-replication/187">An Elegant Experiment to Test the Process of DNA Replication</a></li> <li><a href="/en/library/scientists-and-research/58/the-founding-of-neuroscience/233">The Founding of Neuroscience</a></li> <li><a href="/en/library/scientists-and-research/58/tracking-endangered-jaguars-across-the-border/189">Tracking Endangered Jaguars across the Border</a></li> <li><a href="/en/library/scientists-and-research/58/atmospheric-chemistry-research-that-changed-global-policy/211">Atmospheric Chemistry Research that Changed Global Policy</a></li> <li class="current">Revolutionizing Medicine with Monoclonal Antibodies</li> <li><a href="/en/library/scientists-and-research/58/uncovering-the-mysteries-of-chronic-mountain-sickness/238">Uncovering the Mysteries of Chronic Mountain Sickness</a></li> </ul> </div> <button class="accordion__button" id="acc-button-profiles-in-science" data-accordion="button" aria-controls="acc-panel-profiles-in-science" aria-expanded="false"> <span class="accordion__button__label"> Profiles in Science </span> </button> <div class="accordion__panel" id="acc-panel-profiles-in-science" data-accordion="panel" aria-labelledby="acc-button-profiles-in-science" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/scientists-and-research/58/luis-e.-miramontes/232">Luis E. Miramontes</a></li> <li><a href="/en/library/scientists-and-research/58/bernardo-houssay/237">Bernardo Houssay</a></li> <li><a href="/en/library/scientists-and-research/58/craig-lee/256">Craig Lee</a></li> <li><a href="/en/library/scientists-and-research/58/david-ho/241">David Ho</a></li> <li><a href="/en/library/scientists-and-research/58/louis-tompkins-wright/244">Louis Tompkins Wright</a></li> <li><a href="/en/library/scientists-and-research/58/carlos-j.-finlay/217">Carlos J. Finlay</a></li> <li><a href="/en/library/scientists-and-research/58/cecilia-payne/290">Cecilia Payne</a></li> <li><a href="/en/library/scientists-and-research/58/jazmin-scarlett/291">Jazmin Scarlett</a></li> <li><a href="/en/library/scientists-and-research/58/ramari-stewart/292">Ramari Stewart</a></li> <li><a href="/en/library/scientists-and-research/58/johnson-cerda/300">Johnson Cerda</a></li> <li><a href="/en/library/scientists-and-research/58/ellen-ochoa/201">Ellen Ochoa</a></li> <li><a href="/en/library/scientists-and-research/58/ruth-benerito/205">Ruth Benerito</a></li> <li><a href="/en/library/scientists-and-research/58/franklin-chang-díaz/219">Franklin Chang Díaz</a></li> <li><a href="/en/library/scientists-and-research/58/percy-lavon-julian/221">Percy Lavon Julian</a></li> <li><a href="/en/library/scientists-and-research/58/luis-walter-alvarez/229">Luis Walter Alvarez</a></li> <li><a href="/en/library/scientists-and-research/58/france-anne-dominic-córdova/230">France Anne-Dominic Córdova</a></li> </ul> </div> </div> </div> </div> </div> </li> <li>  <button class="button" data-toggle="dropdown">Scientists and Research </button> <div class="nav__dropdown box-shadow-1 padding-1"> <div class="accordion accordion--secondary font-size-sm"> <div class="accordion accordion--secondary"> <button class="accordion__button" id="acc-sub-button-scientific-research" data-accordion="button" aria-controls="acc-sub-panel-scientific-research" aria-expanded="false"> <span class="accordion__button__label"> Scientific Research </span> </button> <div class="accordion__panel" id="acc-sub-panel-scientific-research" data-accordion="panel" aria-labelledby="acc-sub-button-scientific-research" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/scientists-and-research/58/collaborative-research-in-the-arctic-towards-understanding-climate-change/183">Collaborative Research in the Arctic Towards Understanding Climate Change</a></li> <li><a href="/en/library/scientists-and-research/58/from-stable-chromosomes-to-jumping-genes/184">From Stable Chromosomes to Jumping Genes</a></li> <li><a href="/en/library/scientists-and-research/58/an-elegant-experiment-to-test-the-process-of-dna-replication/187">An Elegant Experiment to Test the Process of DNA Replication</a></li> <li><a href="/en/library/scientists-and-research/58/the-founding-of-neuroscience/233">The Founding of Neuroscience</a></li> <li><a href="/en/library/scientists-and-research/58/tracking-endangered-jaguars-across-the-border/189">Tracking Endangered Jaguars across the Border</a></li> <li><a href="/en/library/scientists-and-research/58/atmospheric-chemistry-research-that-changed-global-policy/211">Atmospheric Chemistry Research that Changed Global Policy</a></li> <li class="current">Revolutionizing Medicine with Monoclonal Antibodies</li> <li><a href="/en/library/scientists-and-research/58/uncovering-the-mysteries-of-chronic-mountain-sickness/238">Uncovering the Mysteries of Chronic Mountain Sickness</a></li> </ul> </div> <button class="accordion__button" id="acc-sub-button-profiles-in-science" data-accordion="button" aria-controls="acc-sub-panel-profiles-in-science" aria-expanded="false"> <span class="accordion__button__label"> Profiles in Science </span> </button> <div class="accordion__panel" id="acc-sub-panel-profiles-in-science" data-accordion="panel" aria-labelledby="acc-sub-button-profiles-in-science" role="region"> <ul class="nav text-color-link"> <li><a href="/en/library/scientists-and-research/58/luis-e.-miramontes/232">Luis E. Miramontes</a></li> <li><a href="/en/library/scientists-and-research/58/bernardo-houssay/237">Bernardo Houssay</a></li> <li><a href="/en/library/scientists-and-research/58/craig-lee/256">Craig Lee</a></li> <li><a href="/en/library/scientists-and-research/58/david-ho/241">David Ho</a></li> <li><a href="/en/library/scientists-and-research/58/louis-tompkins-wright/244">Louis Tompkins Wright</a></li> <li><a href="/en/library/scientists-and-research/58/carlos-j.-finlay/217">Carlos J. Finlay</a></li> <li><a href="/en/library/scientists-and-research/58/cecilia-payne/290">Cecilia Payne</a></li> <li><a href="/en/library/scientists-and-research/58/jazmin-scarlett/291">Jazmin Scarlett</a></li> <li><a href="/en/library/scientists-and-research/58/ramari-stewart/292">Ramari Stewart</a></li> <li><a href="/en/library/scientists-and-research/58/johnson-cerda/300">Johnson Cerda</a></li> <li><a href="/en/library/scientists-and-research/58/ellen-ochoa/201">Ellen Ochoa</a></li> <li><a href="/en/library/scientists-and-research/58/ruth-benerito/205">Ruth Benerito</a></li> <li><a href="/en/library/scientists-and-research/58/franklin-chang-díaz/219">Franklin Chang Díaz</a></li> <li><a href="/en/library/scientists-and-research/58/percy-lavon-julian/221">Percy Lavon Julian</a></li> <li><a href="/en/library/scientists-and-research/58/luis-walter-alvarez/229">Luis Walter Alvarez</a></li> <li><a href="/en/library/scientists-and-research/58/france-anne-dominic-córdova/230">France Anne-Dominic Córdova</a></li> </ul> </div> </div> </div> </div> </li> </ul> </nav>  <div id="theTop"></div> <main id="skip-header-content"> <div class="margin-bottom-5"> <article class="container wide module"> <header class="grid grid--sidebar-right module__header"> <div class="module__header__title"> <span class="subcategory"> <strong><em>Scientific Research</em></strong> </span> <h1>Revolutionizing Medicine with Monoclonal Antibodies: <sub><em>The work of César Milstein</em></sub></h1> <p class="byline">by David Warmflash, MD</p> <nav class="module__header__tabs"> <ul class="tabs-nav tabs-nav--horizontal library"> <li> <a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220/reading" aria-current="page" >Reading</a> </li> <li> <a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220/quiz">Quiz</a> </li> <li> <a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220/resources">Teach with this</a> </li> </ul> </nav> </div> </header> <hr class="divider"/>   <div class="grid grid--sidebar-right grid--divider"> <div class="order-2 order-1--lg module__main"> <div class="narrow margin-x-auto margin-y-5"> <div class="accordion margin-bottom-5">  <button class="accordion__button" id="acc-button-key-concepts" data-accordion="button" aria-controls="acc-panel-key-concepts" aria-expanded="true" tabindex="0"> Did you know? </button> <div class="accordion__panel shown show" id="acc-panel-key-concepts" data-accordion="panel" aria-labelledby="acc-button-key-concepts" role="region"> <div class="accordion__panel__content"> <p>Did you know that cancer cells can be used for a good purpose? In the 1970s, immunologist César Milstein changed medicine by combining two cells: one that was capable of producing disease-fighting antibodies, and the other a cancer cell that enabled the antibody to be reproduced over and over. Virtually every blood test we rely on today to detect specific diseases or even pregnancy uses the technique pioneered by Milstein.</p> </div> </div>  <button class="accordion__button" id="acc-button-terms-you-should-know" data-accordion="button" aria-controls="acc-panel-terms-you-should-know" aria-expanded="false" tabindex="0"> Terms you should know </button> <div class="accordion__panel" id="acc-panel-terms-you-should-know" data-accordion="panel" aria-labelledby="acc-button-terms-you-should-know" role="region" aria-hidden="true"> <div class="accordion__panel__content"> <dl> <dt><a href="/en/glossary/view/antibody">antibody </a></dt> <dd> a Y-shaped protein molecule produced by the body’s immune system that fights disease by binding with a particular antigen </dd> <dt><a href="/en/glossary/view/antigen">antigen </a></dt> <dd> a foreign entity in the body that triggers the body’s immune system to produce antibodies </dd> <dt><a href="/en/glossary/view/mutation">mutation </a></dt> <dd> a change in gene sequences of cell DNA</dd> </dl> </div> </div> </div> <hr class="border-color-dark" /> <section> <div class="container narrow"> <p>On December 8, 1984, immunologist César Milstein arrived at the Karolinska Institutet in Stockholm, Sweden, to accept the <mark class="term" data-term="Nobel Prize" data-term-def="Awards made annually, beginning in 1901, from funds originally established by Alfred B. Nobel for outstanding achievement in physics, chemistry, medicine&hellip;" data-term-url="/en/glossary/view/Nobel+Prize/3843">Nobel Prize</mark> in Physiology and Medicine. Outside the world of immunology, his name is not well-known, but Milstein’s work advanced medicine and science in very profound ways. From routine applications like home pregnancy tests to more exotic jobs like searching for life on Mars, Milstein’s discovery of monoclonal antibodies and their uses has enabled humans to do things that otherwise might not be possible. Beyond tests, monoclonal antibodies are used ever more frequently in medicine – Zmapp, a treatment for Ebola, is just one example.</p> <div class="figure"> <figure> <button class="lightbox-button lightbox-button--icon" data-lightbox="" data-lightbox-src="/img/library/large_images/image_8374.jpg"> <img src="/img/library/modules/mid220/Image/VLObject-8374-150704080718.jpg" alt="Figure 1: César Milstein in the Medical Research Council laboratory." /> </button> <figcaption> <p><strong>Figure 1</strong>: César Milstein in the Medical Research Council laboratory.</p> <span class="credit">image ©MRC Laboratory of Molecular Biology</span> </figcaption> </figure> </div> <p>Monoclonal antibodies come from specially developed <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark> called “hybridomas.” A hybridoma is a combination, or fusion, of two different types of cells: one with the <mark class="term" data-term="gene" data-term-def="Material (usually DNA) that is inherited from a parent and which encodes for a cellular component important for some cellular function." data-term-url="/en/glossary/view/gene/3294">genes</mark> needed to make the desired <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark>, the other a cancer cell that makes the hybridoma “immortal.” This means that the hybridoma can be reproduced over and over to produce cells identical to itself and provide a specific antibody. The idea of a combining a cancer cell with another cell for a good purpose may sound counterintuitive, but in the 1970s Milstein and his colleagues realized the enormous potential of this technology to change medicine and biology. </p> <p><section id="toc_1" class=""> <h2>Snake venom as inspiration</h2></p> <p>Born October 8, 1927 in Bahía Blanca, Argentina, César Milstein was the middle of three brothers in a family of Jewish immigrants from Eastern Europe. His father, Lazaro, hailed from a village in Ukraine; his mother Maxima was born on Argentinian <mark class="term" data-term="soil" data-term-def="The loose top layer of Earth’s surface where plants grow, made up of particles of rocks, minerals, and organic material." data-term-url="/en/glossary/view/soil/8563">soil</mark>, but she herself was a child of Ukrainian Jews. Like many East European Jews of that time <mark class="term" data-term="period" data-term-def="A row of elements in the periodic table." data-term-url="/en/glossary/view/period/8565">period</mark>, Milstein’s parents identified as Jews strongly in the cultural sense, and not at all in the religious sense. They read Yiddish literature and associated socially with other Jews, which included working in non-religious Jewish organizations. They were part of socialist movements, concerned with workers’ rights, and did not attend synagogue or view the world from a religious perspective. They also spoke Yiddish with one another at home, but raised their sons to speak only Spanish.</p><p>As a child, Milstein loved reading books and became fascinated with science at age 8 because of a conversation with an older cousin. The cousin worked at the Instituto Malbran as a biochemist, and during their conversation she explained to the young César how she was developing <mark class="term" data-term="serum" data-term-def="(Plural: sera) The liquid portion of whole blood after other material (cells and clotting factors) are removed. Blood serum does contain&hellip;" data-term-url="/en/glossary/view/serum/10459">serum</mark> from snake bite victims to be used as a snake bite treatment. This was the beginning of Milstein’s fascination with the immune system, and it had a major impact on his future work. Back then, scientists did not know the mechanism underlying the effect of a snake bite antiserum. But later, Milstein learned that the immune system produced antibodies, <mark class="term" data-term="protein" data-term-def="Macromolecules that are polymers of individual amino acids arranged in a chain and joined together by peptide bonds (and so also&hellip;" data-term-url="/en/glossary/view/protein/1594">proteins</mark> that gave the antiserum its beneficial effect against the toxins that people received in a snake bite.</p><p>A year after the conversation with his cousin about the snake <mark class="term" data-term="serum" data-term-def="(Plural: sera) The liquid portion of whole blood after other material (cells and clotting factors) are removed. Blood serum does contain&hellip;" data-term-url="/en/glossary/view/serum/10459">serum</mark>, Milstein read a Spanish translation of <em>Microbe Hunters</em> by Paul de Kruif. The book described Antony van Leeuwenhoek, <mark class="term" data-term="Louis Pasteur" data-term-def="A French chemist and biologist, born in Dole, France (1822-1895). Pasteur founded the science of microbiology and proved that microorganisms cause&hellip;" data-term-url="/en/glossary/view/Pasteur%2C+Louis/4468">Louis Pasteur</mark>, and other pioneers of biology as adventurers. This drew Milstein further into the world of life science. He knew at that point that he wanted a biological career. </p> <div class="figure"> <figure> <button class="lightbox-button lightbox-button--icon" data-lightbox=""> <img src="/img/library/modules/mid220/Image/VLObject-8375-150704080741.jpg" alt="Figure 2: Los Cazadores de Microbios, the Spanish version of The Microbe Hunters by Dr. Paul de Kruif." /> </button> <figcaption> <p><strong>Figure 2</strong>: <em>Los Cazadores de Microbios</em>, the Spanish version of <em>The Microbe Hunters</em> by Dr. Paul de Kruif.</p> </figcaption> </figure> </div> </section> <section id="toc_2"> <h2>A student and activist in Buenos Aires</h2><p>Milstein moved to Argentina’s capital, Buenos Aires, for his high school education. In 1945, he entered the University of Buenos Aires as a chemistry major. He was a brilliant science student, but politics and funding shortages forced him to think about other matters alongside his studies. His secular Jewish, left-wing upbringing lay at the foundation of his identity and his interest in science did not replace that, but added to it. As an undergraduate, he naturally gravitated into student movements against the Perón government that ruled Argentina. This government was right-wing economically and generally did not support scientific research at the university. On top of that, the right-wing government was complex when it came to relations with the country’s Jewish community. The founder of the rightwing party, Juan Perón, sympathized with the Axis powers of World War II, and Nazis fleeing Europe were allowed refuge in Argentina. On the other hand, Perón spoke in favor of Jewish rights and established diplomatic relations with the state of Israel early in 1949, and there were noticeable numbers of Jewish Peronists. But Milstein’s Jewish family was socialist. He was raised to believe that lack of wealth should not be a barrier to the benefits of civilization, especially education.</p> <div class="figure"> <figure> <button class="lightbox-button lightbox-button--icon" data-lightbox=""> <img src="/img/library/modules/mid220/Image/VLObject-8377-150704090712.jpg" alt="Figure 3: A chemistry lab at the University of Buenos Aires in 1947." /> </button> <figcaption> <p><strong>Figure 3</strong>: A chemistry lab at the University of Buenos Aires in 1947.</p> <span class="credit">image ©Digital Library / Program FCEN History, Faculty of Natural Sciences, University of Buenos Aires</span> </figcaption> </figure> </div> <p>The right-wing Peronist policies aggravated most students because they put restrictions on universities and student life and sought to privatize education. Privatizing would mean that only students from wealthy families would be able to attend college. Milstein was popular on campus because he sided with left-wing student movements that favored free education, and in 1951 he ended up as president of the student union. It was a big risk for Milstein because student leaders were being arrested and the student union president was expected to help them. The most famous arrested student was Ernesto Mario Bravo. Like Milstein, Bravo was a chemistry student. On May 17, 1951, Bravo was abducted by the police (not actually arrested) and tortured by the government for the next 20 days. Students and university administrators protested, demanding Bravo’s return, and the protests culminated in a two-day university strike. As student union president, Milstein was under the government spotlight but the protests and strike led to Bravo’s release.</p><p>Along with intense campus politics, Milstein’s undergraduate years were difficult because of an accident that he suffered from diving into a pond and hitting a log. To recover, he had to take some time off. After returning to school, he fell in love with Celia Prilleltensky, a fellow chemistry student. The two students graduated in 1951 and were married one year later. At that time, Milstein enrolled as a graduate student at the same university and found a faculty advisor, Professor Andrés Stoppani, to guide him through a PhD program in biochemistry. When he began the PhD program in 1951, Milstein was shocked and disappointed to learn how underfunded Stoppani and his research lab were. As with the issues that Milstein had faced as an undergraduate, the underfunding of research was the result of the Peronist right-wing policies. Stoppani suspected that Milstein’s political perspective, and his history campaigning against the Peronist education policy, would get the young man into trouble. Stoppani advised Milstein to take time off with his wife until the political environment changed. </p><p>By 1954, the political situation had calmed enough for Milstein to start working with Stoppani. At the time, Stoppani still had no funding to support a graduate student. He was forced to use a portion of his very low salary to acquire materials for <mark class="term" data-term="experiment" data-term-def="A test or trial carried out under controlled conditions so that specific actions can be performed and the results can be observed." data-term-url="/en/glossary/view/experiment/8292">experiments</mark> and he could not afford needed equipment. Milstein worked on <mark class="term" data-term="enzyme" data-term-def="Molecules produced by living organisms that help catalyze biochemical reactions. Enzymes are predominantly protein or protein-based molecules and are highly&hellip;" data-term-url="/en/glossary/view/enzyme/1595">enzyme</mark> research in the Stoppani lab for his dissertation. However, to support his studies, he had to work part-time in a private clinical biochemistry laboratory. Only in 1955 did funding from the government improve enough for the biochemistry department to buy some basic equipment, such as a refrigerated centrifuge, but other essential machines were still out of grasp. To use one particularly important machine – a spectrophotometer – Milstein had to walk several blocks between buildings. This used up a lot of precious time and one day also caused him to break some expensive glassware, which almost got him expelled from the department.</p><p>Milstein’s devotion to politics on campus as an undergraduate and the lack of funding that plagued his day-to-day life as a graduate student drew time away from his studies. Nevertheless, he performed brilliantly as a student. In 1957, he earned his doctorate based on research concerning a type of chemical <mark class="term" data-term="bond" data-term-def="The force that holds together units such as atoms or molecules. <br> <b>[verb]</b> To hold or fasten units such as atoms or molecules together." data-term-url="/en/glossary/view/bond/8297">bond</mark> in <mark class="term" data-term="enzyme" data-term-def="Molecules produced by living organisms that help catalyze biochemical reactions. Enzymes are predominantly protein or protein-based molecules and are highly&hellip;" data-term-url="/en/glossary/view/enzyme/1595">enzymes</mark> called a disulfide bridge. Along with earning his PhD, the disulfide bridge research won him an Argentinian Chemical Association award for best thesis. From that point on, he continued working with Stoppani and the two published groundbreaking scientific papers resulting from Milstein’s doctoral work.</p> <div class="comprehension-checkpoint margin-y-4"> <h6 class="comprehension-checkpoint__header"> <span> <span class="icon icon-question"></span> </span> Comprehension Checkpoint </h6> <form class="" name="cc8342"> <div class="form-entry"> <div class="form-entry__field"> <span class="form-entry__field__label">In Argentina, science education was well-funded during the Peron regime.</span> <div class="form-entry__option"> <div class="form-entry__option__radio" data-answer="incorrect"> <label> <input id="q1-8342-0-option-a" name="quiz-option-8342" type="radio" value="true" > <span class="option__label"> <span class="screen-reader-only">a.</span> true </span> </label> <span class="quiz__response" id="response-8342-0"> <strong>Incorrect.</strong> </span> </div> <div class="form-entry__option__radio" data-answer="correct"> <label> <input id="q1-8342-1-option-b" name="quiz-option-8342" type="radio" value="false" > <span class="option__label"> <span class="screen-reader-only">b.</span> false </span> </label> <span class="quiz__response" id="response-8342-1"> <strong>Correct!</strong> </span> </div> </div> </div> </div> </form> </div> </section> <section id="toc_3"> <h2>Professional advances, but more political difficulties</h2><p>In 1958, Milstein moved to Cambridge, England to continue his research at the Sir William Dunn School of Biochemistry, supported by a fellowship from the British Council. Within a couple of years, this led Milstein to be awarded a second doctorate, this one from Cambridge University, based on his research of an <mark class="term" data-term="enzyme" data-term-def="Molecules produced by living organisms that help catalyze biochemical reactions. Enzymes are predominantly protein or protein-based molecules and are highly&hellip;" data-term-url="/en/glossary/view/enzyme/1595">enzyme</mark> called <em>phosphoglucomutase</em> – research that revealed a very unexpected mechanism through which the enzyme is activated. During this <mark class="term" data-term="period" data-term-def="A row of elements in the periodic table." data-term-url="/en/glossary/view/period/8565">period</mark>, Milstein also met and formed professional bonds with the famed biochemist Fred Sanger, winner of the 1958 <mark class="term" data-term="Nobel Prize" data-term-def="Awards made annually, beginning in 1901, from funds originally established by Alfred B. Nobel for outstanding achievement in physics, chemistry, medicine&hellip;" data-term-url="/en/glossary/view/Nobel+Prize/3843">Nobel Prize</mark>. Soon, Milstein and Sanger started working together on the phosphoglucomutase enzyme. In 1960, the pair published a paper revealing the sequence of amino <mark class="term" data-term="acid" data-term-def="Generally, a substance that reacts with bases to form a salt, several different definitions of acids have been proposed by different&hellip;" data-term-url="/en/glossary/view/acid/1573">acids</mark> (building blocks of proteins) that made up an important region of the enzyme. </p> <div class="figure"> <figure> <button class="lightbox-button lightbox-button--icon" data-lightbox="" data-lightbox-src="/img/library/large_images/image_8378.jpg"> <img src="/img/library/modules/mid220/Image/VLObject-8378-150704090745.jpg" alt="Figure 4: César Milstein and Fred Sanger" /> </button> <figcaption> <p><strong>Figure 4</strong>: César Milstein and Fred Sanger</p> <span class="credit">image ©MRC Laboratory of Molecular Biology</span> </figcaption> </figure> </div> <p>In 1961, Milstein returned to Argentina where he became head of the new Department of Molecular Biology at the Instituto Malbran, the same research center where his older cousin had worked on snake antiserum. His wife Celia also was appointed to the new department. It was a time of reform, with many other scientists returning to Argentina alongside the Milsteins, because the Perón government had fallen. In addition to continuing to research phosphoglucomutase, Milstein started studying another <mark class="term" data-term="enzyme" data-term-def="Molecules produced by living organisms that help catalyze biochemical reactions. Enzymes are predominantly protein or protein-based molecules and are highly&hellip;" data-term-url="/en/glossary/view/enzyme/1595">enzyme</mark>, <em>alkaline phosphatase</em>. Very soon, however, the new government was overthrown by a coup d'etat. This put in place another right-wing government, one hostile to Milstein because he was an academic and his Jewish name, which in the mind of the authorities made him a suspected communist. And so, along with thousands of other scientists and academics, Milstein left Argentina once again and returned to Cambridge.</p> <div class="comprehension-checkpoint margin-y-4"> <h6 class="comprehension-checkpoint__header"> <span> <span class="icon icon-question"></span> </span> Comprehension Checkpoint </h6> <form class="" name="cc8346"> <div class="form-entry"> <div class="form-entry__field"> <span class="form-entry__field__label">Milstein focused his research primarily on</span> <div class="form-entry__option"> <div class="form-entry__option__radio" data-answer="correct"> <label> <input id="q1-8346-0-option-a" name="quiz-option-8346" type="radio" value="enzymes." > <span class="option__label"> <span class="screen-reader-only">a.</span> enzymes. </span> </label> <span class="quiz__response" id="response-8346-0"> <strong>Correct!</strong> </span> </div> <div class="form-entry__option__radio" data-answer="incorrect"> <label> <input id="q1-8346-1-option-b" name="quiz-option-8346" type="radio" value="venomous snakes." > <span class="option__label"> <span class="screen-reader-only">b.</span> venomous snakes. </span> </label> <span class="quiz__response" id="response-8346-1"> <strong>Incorrect.</strong> </span> </div> </div> </div> </div> </form> </div> </section> <section id="toc_4"> <h2>Antibody research</h2><p>Milstein was able to move back to Cambridge quickly because of his friend and colleague Fred Sanger, who directed the <mark class="term" data-term="protein" data-term-def="Macromolecules that are polymers of individual amino acids arranged in a chain and joined together by peptide bonds (and so also&hellip;" data-term-url="/en/glossary/view/protein/1594">protein</mark> chemistry division of the Medical Research Council Laboratory of Molecular Biology (LMB). Milstein’s focus at LMB was on the formation of <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark> <mark class="term" data-term="molecule" data-term-def="A particle formed by the chemical bonding of two or more atoms. The molecule is the smallest particle of a&hellip;" data-term-url="/en/glossary/view/molecule/1518">molecules</mark>, working on the very topic that had inspired him as a boy. </p> <div class="figure"> <figure> <button class="lightbox-button lightbox-button--icon" data-lightbox="" data-lightbox-src="/img/library/large_images/image_8380.jpg"> <img src="/img/library/modules/mid220/Image/VLObject-8380-150704100732.jpg" alt="Figure 5: Schematic diagram of an antibody and antigens with light and heavy chains noted." /> </button> <figcaption> <p><strong>Figure 5</strong>: Schematic diagram of an antibody and antigens with light and heavy chains noted.</p> </figcaption> </figure> </div> <p>Scientists had known about antibodies since the late 19<sup>th</sup> century. They knew that antibodies protected against disease, but they had no idea how antibodies worked. By the time Milstein started his research at LMB in the 1960s, scientists knew that antibodies were <mark class="term" data-term="protein" data-term-def="Macromolecules that are polymers of individual amino acids arranged in a chain and joined together by peptide bonds (and so also&hellip;" data-term-url="/en/glossary/view/protein/1594">proteins</mark>. They knew that antibodies were shaped almost like the letter <em>Y</em> and that this shape had something to do with how antibodies worked. The top part of the <em>Y</em> shape was known to vary enormously between antibodies. This variability allows the immune system to produce antibodies able to grab onto a wide range of <mark class="term" data-term="molecule" data-term-def="A particle formed by the chemical bonding of two or more atoms. The molecule is the smallest particle of a&hellip;" data-term-url="/en/glossary/view/molecule/1518">molecules</mark> foreign to the body. The upper part of the <em>Y</em> shape grabs a particular foreign molecule the way that a lock connects perfectly with the shape of the key that matches it. Scientists in the 1960s knew that antibodies worked this way, but how so many different shapes could be produced was a mystery.</p><p>To solve the mystery, Milstein designed <mark class="term" data-term="experiment" data-term-def="A test or trial carried out under controlled conditions so that specific actions can be performed and the results can be observed." data-term-url="/en/glossary/view/experiment/8292">experiments</mark> to test the possibility that the diversity could be the result of <mark class="term" data-term="mutation" data-term-def="A change in gene sequences of cell DNA." data-term-url="/en/glossary/view/mutation/8757">mutations</mark> occurring in the <mark class="term" data-term="DNA" data-term-def="Deoxyribonucleic acid. A double-stranded nucleic acid containing the sugar 2-deoxy-D-ribose. A constituent of cellular nuclear material responsible for encoding&hellip;" data-term-url="/en/glossary/view/DNA/1604">DNA</mark> sequences. This <mark class="term" data-term="hypothesis" data-term-def="From the Greek word <em>hypothesis</em> meaning assumption or the basis of an argument, a hypothesis is a proposal intended to explain&hellip;" data-term-url="/en/glossary/view/hypothesis/3727">hypothesis</mark> was developed based on an earlier, simpler idea proposed in 1959 by another pioneering molecular biologist, Joshua Lederberg. Milstein was not talking about the kind of mutations that are passed on to future <mark class="term" data-term="generation" data-term-def="Offspring at the same step in the line of descent from a common ancestor." data-term-url="/en/glossary/view/generation/8293">generations</mark>. Instead, he meant <em>somatic mutations</em>, changes in <mark class="term" data-term="gene" data-term-def="Material (usually DNA) that is inherited from a parent and which encodes for a cellular component important for some cellular function." data-term-url="/en/glossary/view/gene/3294">gene</mark> sequences in the DNA of body <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark>, which are all cells other than reproductive cells (called gametes). These mutations not passed on to the next generation, but are important because they often lead to cancer. In particular, Milstein set out to investigate somatic mutations in the immune system cells that make antibodies.</p> <div class="comprehension-checkpoint margin-y-4"> <h6 class="comprehension-checkpoint__header"> <span> <span class="icon icon-question"></span> </span> Comprehension Checkpoint </h6> <form class="" name="cc8351"> <div class="form-entry"> <div class="form-entry__field"> <span class="form-entry__field__label">In antibodies, the top of their <em>Y</em>-shape</span> <div class="form-entry__option"> <div class="form-entry__option__radio" data-answer="incorrect"> <label> <input id="q1-8351-0-option-a" name="quiz-option-8351" type="radio" value="is the same in every antibody." > <span class="option__label"> <span class="screen-reader-only">a.</span> is the same in every antibody. </span> </label> <span class="quiz__response" id="response-8351-0"> <strong>Incorrect.</strong> </span> </div> <div class="form-entry__option__radio" data-answer="correct"> <label> <input id="q1-8351-1-option-b" name="quiz-option-8351" type="radio" value="varies enormously among different antibodies." > <span class="option__label"> <span class="screen-reader-only">b.</span> varies enormously among different antibodies. </span> </label> <span class="quiz__response" id="response-8351-1"> <strong>Correct!</strong> </span> </div> </div> </div> </div> </form> </div> </section> <section id="toc2_1"><h3>Tackling research problems</h3><p>During the 1960s, numerous scientists were studying the <mark class="term" data-term="DNA" data-term-def="Deoxyribonucleic acid. A double-stranded nucleic acid containing the sugar 2-deoxy-D-ribose. A constituent of cellular nuclear material responsible for encoding&hellip;" data-term-url="/en/glossary/view/DNA/1604">DNA</mark> of immune system <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark> and they all, including Milstein, encountered the same two problems. First, <mark class="term" data-term="experiment" data-term-def="A test or trial carried out under controlled conditions so that specific actions can be performed and the results can be observed." data-term-url="/en/glossary/view/experiment/8292">experiments</mark> required a large number of cells and cells that made antibodies were hard to isolate. Second, the research required figuring out how the DNA in a cell that made one type of <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark> differed from the DNA of a cell that made a different antibody. Every person has billions of antibody-producing immune cells in his or her blood. Each cell that is dedicated to making a specific antibody is called a clone. To compare one clone with another, Milstein knew he would need multiple copies of each clone.</p><p>To overcome both problems, Milstein investigated <em>Bence-Jones proteins</em>, which are produced by a type of cancer called multiple myeloma. Antibodies are composed of smaller units that form the upper part of the <em>Y</em>-shape called heavy chains and light chains. Since the Bence-Jones <mark class="term" data-term="protein" data-term-def="Macromolecules that are polymers of individual amino acids arranged in a chain and joined together by peptide bonds (and so also&hellip;" data-term-url="/en/glossary/view/protein/1594">proteins</mark> appeared to have the same structure as a section of the light chains of antibodies, this solved the problem of isolating antibodies. The Bence-Jones proteins could also be obtained in large quantities from the urine and blood of multiple myeloma patients, solving the problem of acquiring large numbers of <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark>. Furthermore, Bence-Jones protein <mark class="term" data-term="molecule" data-term-def="A particle formed by the chemical bonding of two or more atoms. The molecule is the smallest particle of a&hellip;" data-term-url="/en/glossary/view/molecule/1518">molecules</mark> from the same patient were known to be identical to one another. Essentially, multiple myeloma makes a whole bunch of the same immune cell clone and a piece of that <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark>, the light chain, accumulates in the person’s blood and urine. </p> <div class="figure"> <figure> <button class="lightbox-button lightbox-button--icon" data-lightbox="" data-lightbox-src="/img/library/large_images/image_8381.jpg"> <img src="/img/library/modules/mid220/Image/VLObject-8381-150705070747.jpg" alt="Figure 6: A crystal of Bence-Jones protein created with X-ray crystallography, which can reveal detailed, three-dimensional protein structures." /> </button> <figcaption> <p><strong>Figure 6</strong>: A crystal of Bence-Jones protein created with X-ray crystallography, which can reveal detailed, three-dimensional protein structures.</p> <span class="credit">image ©NIH / Alex McPherson, University of California, Irvine</span> </figcaption> </figure> </div> <p>By utilizing Bence-Jones <mark class="term" data-term="protein" data-term-def="Macromolecules that are polymers of individual amino acids arranged in a chain and joined together by peptide bonds (and so also&hellip;" data-term-url="/en/glossary/view/protein/1594">proteins</mark> from patient samples, Milstein believed he might figure out faster than other scientists how antibodies worked and how they were formed. The hope was that it would lead him to devise <mark class="term" data-term="experiment" data-term-def="A test or trial carried out under controlled conditions so that specific actions can be performed and the results can be observed." data-term-url="/en/glossary/view/experiment/8292">experiments</mark> to test his somatic <mark class="term" data-term="mutation" data-term-def="A change in gene sequences of cell DNA." data-term-url="/en/glossary/view/mutation/8757">mutation</mark> <mark class="term" data-term="hypothesis" data-term-def="From the Greek word <em>hypothesis</em> meaning assumption or the basis of an argument, a hypothesis is a proposal intended to explain&hellip;" data-term-url="/en/glossary/view/hypothesis/3727">hypothesis</mark>, that the diversity of <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark> shapes is the result of mutations in the <mark class="term" data-term="DNA" data-term-def="Deoxyribonucleic acid. A double-stranded nucleic acid containing the sugar 2-deoxy-D-ribose. A constituent of cellular nuclear material responsible for encoding&hellip;" data-term-url="/en/glossary/view/DNA/1604">DNA</mark> sequences. Like the protein <mark class="term" data-term="enzyme" data-term-def="Molecules produced by living organisms that help catalyze biochemical reactions. Enzymes are predominantly protein or protein-based molecules and are highly&hellip;" data-term-url="/en/glossary/view/enzyme/1595">enzymes</mark> that Milstein had studied, antibodies and Bence-Jones proteins are made of chemical building blocks called amino <mark class="term" data-term="acid" data-term-def="Generally, a substance that reacts with bases to form a salt, several different definitions of acids have been proposed by different&hellip;" data-term-url="/en/glossary/view/acid/1573">acids</mark>. The sequence of amino acids of each protein depends on DNA sequences, or <mark class="term" data-term="gene" data-term-def="Material (usually DNA) that is inherited from a parent and which encodes for a cellular component important for some cellular function." data-term-url="/en/glossary/view/gene/3294">genes</mark>, available in each <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cell</mark>. Milstein’s long-term tactic was to compare the sequences of DNA and amino acids with the structure of the Bence-Jones protein. If he could do this, he could prove that the diversity of antibodies does indeed come from somatic mutations.</p><p>In 1962, Michael Potter, a molecular biologist at the US National Cancer Institute (NCI), discovered accidentally that a certain <mark class="term" data-term="strain" data-term-def="A group of closely related organisms; a distinct variety, as in a strain of bacteria." data-term-url="/en/glossary/view/strain/8288">strain</mark> of laboratory mice (BALB/c mice) grows myeloma <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark> if injected with mineral oil. This made multiple myeloma cells and Bence-Jones <mark class="term" data-term="protein" data-term-def="Macromolecules that are polymers of individual amino acids arranged in a chain and joined together by peptide bonds (and so also&hellip;" data-term-url="/en/glossary/view/protein/1594">proteins</mark> much easier to obtain compared with getting them from the blood and urine of human volunteers. Potter and the scientists working with him at NCI started growing large supplies of myeloma cells and made them available to scientists around the world. During the 1970s a team at San Diego’s Salk Institute developed a way to grow Potter’s cells in tissue culture; this meant that large numbers of the cells could be grown without even using mice. One line of these cells was called <em>MOPC21</em> and it gave scientists like Milstein the freedom to focus their time and attention on doing creative <mark class="term" data-term="experiment" data-term-def="A test or trial carried out under controlled conditions so that specific actions can be performed and the results can be observed." data-term-url="/en/glossary/view/experiment/8292">experiments</mark> rather than having to devote much of their time into caring for the cells and keeping them alive in mice. Milstein collaborated with pathologist George Brownlee to extract a <mark class="term" data-term="molecule" data-term-def="A particle formed by the chemical bonding of two or more atoms. The molecule is the smallest particle of a&hellip;" data-term-url="/en/glossary/view/molecule/1518">molecule</mark> called <mark class="term" data-term="RNA" data-term-def="Ribonucleic acid. A single-stranded nucleic acid containing the sugar ribose. In most organisms, a molecule responsible for transfer of the&hellip;" data-term-url="/en/glossary/view/RNA/1605">RNA</mark> from the MOPC21 cells. RNA is produced from <mark class="term" data-term="DNA" data-term-def="Deoxyribonucleic acid. A double-stranded nucleic acid containing the sugar 2-deoxy-D-ribose. A constituent of cellular nuclear material responsible for encoding&hellip;" data-term-url="/en/glossary/view/DNA/1604">DNA</mark> and acts as an intermediate in the construction of a protein, including an <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark>, from amino <mark class="term" data-term="acid" data-term-def="Generally, a substance that reacts with bases to form a salt, several different definitions of acids have been proposed by different&hellip;" data-term-url="/en/glossary/view/acid/1573">acid</mark> building blocks. So, by examining the cells’ RNA, the genetic sequence for antibodies made by the cell could be obtained.</p> <div class="comprehension-checkpoint margin-y-4"> <h6 class="comprehension-checkpoint__header"> <span> <span class="icon icon-question"></span> </span> Comprehension Checkpoint </h6> <form class="" name="cc8357"> <div class="form-entry"> <div class="form-entry__field"> <span class="form-entry__field__label">Every person has ______ antibody-producing immune cells in his or her blood.</span> <div class="form-entry__option"> <div class="form-entry__option__radio" data-answer="correct"> <label> <input id="q1-8357-0-option-a" name="quiz-option-8357" type="radio" value="billions of" > <span class="option__label"> <span class="screen-reader-only">a.</span> billions of </span> </label> <span class="quiz__response" id="response-8357-0"> <strong>Correct!</strong> </span> </div> <div class="form-entry__option__radio" data-answer="incorrect"> <label> <input id="q1-8357-1-option-b" name="quiz-option-8357" type="radio" value="very few" > <span class="option__label"> <span class="screen-reader-only">b.</span> very few </span> </label> <span class="quiz__response" id="response-8357-1"> <strong>Incorrect.</strong> </span> </div> </div> </div> </div> </form> </div> </section> <section id="toc2_2"><h3>Searching for mutations</h3><p>In the early 1970s Milstein also began working with two younger scientists, David Secher and Dick Cotton. The research that took shape consisted of two major components. The first was to clone the MOPC21 <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark> to reproduce numerous exact genetic copies of the cell line. These needed to grow on a substance called “soft agar” because this would allow individual clones to be sampled easily. </p><p>The team needed to determine the rate at which <mark class="term" data-term="mutation" data-term-def="A change in gene sequences of cell DNA." data-term-url="/en/glossary/view/mutation/8757">mutations</mark> occurred once clones were growing consistently. Clones that mutated from a <mark class="term" data-term="parent" data-term-def="The material or source from which something is derived." data-term-url="/en/glossary/view/parent/1618">parent</mark> clone – called <em>variants</em> – could then be pulled from the agar, and the effects on the antibodies made by the <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark> could be compared. But after three months of culturing and analyzing antibodies from 7,000 clones, only five variants in <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark> structure were observed. The method was too slow, although the occurrence of any mutations was a good finding.</p></section> <section id="toc_5"> <h2>A new type of cell: Hybridomas</h2><p>The second component of the research sought to fuse two <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark>, each from a different myeloma cell line. This project depended largely on the laboratory skills of Dick Cotton, who was a postdoctoral scientist visiting Milstein’s LMB lab from Australia. All body cells have a mix of <mark class="term" data-term="gene" data-term-def="Material (usually DNA) that is inherited from a parent and which encodes for a cellular component important for some cellular function." data-term-url="/en/glossary/view/gene/3294">genes</mark>; half from the individual’s mother and half from the father. By the 1970s, however, scientists knew that each antibody-making immune cell used just one of the two sets of parental genes to make its <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark>. Either the <mark class="term" data-term="maternal" data-term-def="Derived from or related to one's mother." data-term-url="/en/glossary/view/maternal/5315">maternal</mark> genes were used or the <mark class="term" data-term="paternal" data-term-def="Derived from or related to one's father." data-term-url="/en/glossary/view/paternal/8364">paternal</mark> genes, while the other set of genes was turned off. </p><p>Cotton wanted to understand why this was the case, and together with Milstein he decided that merging two <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark> into one might reveal something about which set of <mark class="term" data-term="gene" data-term-def="Material (usually DNA) that is inherited from a parent and which encodes for a cellular component important for some cellular function." data-term-url="/en/glossary/view/gene/3294">genes</mark> turns on or off in different situations. In the resulting fusion or <mark class="term" data-term="hybrid" data-term-def="Pertaining to the offspring of two plants or animals of different breeds, varieties, species, or genera. Hybrid often refers to plants&hellip;" data-term-url="/en/glossary/view/hybrid/5314">hybrid</mark> cell, they wanted to see which genes would be turned on and which would be turned off. They also wanted to see what effect the turning on or off of different genes in the fused cell would have on the structure of the antibodies made by the hybrid cell. </p><p>For one of the <mark class="term" data-term="parent" data-term-def="The material or source from which something is derived." data-term-url="/en/glossary/view/parent/1618">parent</mark> <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark> to make the <mark class="term" data-term="hybrid" data-term-def="Pertaining to the offspring of two plants or animals of different breeds, varieties, species, or genera. Hybrid often refers to plants&hellip;" data-term-url="/en/glossary/view/hybrid/5314">hybrid</mark> cell, they used a <mark class="term" data-term="mutant" data-term-def="An organism that has undergone a change in genetic material." data-term-url="/en/glossary/view/mutant/8739">mutant</mark> grown from the mouse MOPC21 cell line, while the other parent cell was from a line developed by a Belgian researcher. But rather than showing <mark class="term" data-term="gene" data-term-def="Material (usually DNA) that is inherited from a parent and which encodes for a cellular component important for some cellular function." data-term-url="/en/glossary/view/gene/3294">genes</mark> from one parent turned on and genes from the other parent turned off, the cell made by fusing the mouse and rat cell did something surprising. It manufactured antibodies using genes from both parent cells. Unlike in conventional immune cells, the new hybrid cell, which they called a <em>hybridoma</em>, did not shuffle the genes for different regions of the <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark> to make new kinds of antibodies. Instead, they decided that the gene shuffling must occur early in the development of an immune cell before the cell starts making antibodies.</p><p>Using the same technique, but tweaking procedures to perfect it, the team made other types of mouse-rat hybridomas and also mouse-mouse hybridomas. The results were the same as in the first hybridoma: <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark> <mark class="term" data-term="gene" data-term-def="Material (usually DNA) that is inherited from a parent and which encodes for a cellular component important for some cellular function." data-term-url="/en/glossary/view/gene/3294">genes</mark> from both <mark class="term" data-term="parent" data-term-def="The material or source from which something is derived." data-term-url="/en/glossary/view/parent/1618">parents</mark> were used to make antibodies. There was a problem, however. The new hybridomas revealed a lot about the genetics of antibody production, but there was no way to control which antibody was produced. In normal physiology, the immune system makes antibodies that specifically recognize and bind with a particular foreign entity called an <em>antigen</em>. Since the hybridomas were made from myeloma <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark>, the antibodies produced were the same antibodies that the myeloma cells were born to make as cancer cells. Milstein needed a way to trigger the hybridomas to make new antibodies, meaning antibodies against a particular <mark class="term" data-term="antigen" data-term-def="A substance that stimulates the production of an antibody by the immune system. Antigens include toxins, bacteria, foreign blood cells, and&hellip;" data-term-url="/en/glossary/view/antigen/5280">antigen</mark>, just like the immune system does. Also, the hybridomas were very short lived, which made continuous <mark class="term" data-term="experiment" data-term-def="A test or trial carried out under controlled conditions so that specific actions can be performed and the results can be observed." data-term-url="/en/glossary/view/experiment/8292">experiments</mark> difficult over time.</p><p>While presenting a paper in Basel, Switzerland, Milstein met another researcher: Georges Köhler. For about a decade, Köhler had been working on his own kind of fusion <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark> using normal B <mark class="term" data-term="lymphocyte" data-term-def="Second most abundant type of white blood cell in the blood of vertebrates." data-term-url="/en/glossary/view/lymphocyte/10186">lymphocytes</mark>. Also called <em>B-cells</em>, these are immune system cells that normally make antibodies in response to an infection. When this happens, they’re called <em>plasma cells</em>; when plasma cells become cancerous, they become multiple myeloma, the kind of cells that Milstein was fusing into hybridomas. A new collaboration developed as a result of this meeting and after many <mark class="term" data-term="experiment" data-term-def="A test or trial carried out under controlled conditions so that specific actions can be performed and the results can be observed." data-term-url="/en/glossary/view/experiment/8292">experiments</mark> Milstein and his colleagues created a new kind of hybridoma: a fusion of a healthy human B-cell and a mouse myeloma cell. They did this through a trial and error <mark class="term" data-term="process" data-term-def="Method, procedure; series of actions or steps." data-term-url="/en/glossary/view/process/8256">process</mark>, trying different combinations of cells and observing what the different, resulting hybridomas could do. The human B cell/mouse myeloma hybridoma turned out to be a groundbreaking achievement. It could manufacture the <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark> that normally could be made by its human B-cell <mark class="term" data-term="parent" data-term-def="The material or source from which something is derived." data-term-url="/en/glossary/view/parent/1618">parent</mark>, but, somehow, merging with the mouse myeloma cell made it immortal. This meant that a hybridoma could be used to generate a cell line that could exist for long <mark class="term" data-term="period" data-term-def="A row of elements in the periodic table." data-term-url="/en/glossary/view/period/8565">periods</mark> of time and be used to create the antibody of the immune cell in large quantities.</p> <div class="figure"> <figure> <button class="lightbox-button lightbox-button--icon" data-lightbox="" data-lightbox-src="/img/library/large_images/image_8383.jpg"> <img src="/img/library/modules/mid220/Image/VLObject-8383-150705080734.jpg" alt="Figure 7: An illustration of the steps in creating the human B cell/mouse myeloma hybridoma. (1) Mouse immunized; (2) B-cells isolated from the spleen; (3) Myeloma cells cultivated; (4) Myeloma and B-cells fused; (5) Cell lines separated; (6) Cell lines screened into those that bind with specific antigens (a) and those that do not (b); (7) Multiplication of cells in vitro (a) or in vivo (b); (8) Antibodies harvested" /> </button> <figcaption> <p><strong>Figure 7</strong>: An illustration of the steps in creating the human B cell/mouse myeloma hybridoma. (1) Mouse immunized; (2) B-cells isolated from the spleen; (3) Myeloma cells cultivated; (4) Myeloma and B-cells fused; (5) Cell lines separated; (6) Cell lines screened into those that bind with specific antigens (a) and those that do not (b); (7) Multiplication of cells in vitro (a) or in vivo (b); (8) Antibodies harvested</p> <span class="credit">image ©Martin Brändli</span> </figcaption> </figure> </div> <p>This achievement set up Milstein, Köhler, and a growing number of scientific colleagues for the next step: making a hybridoma that secretes antibodies against a chosen <mark class="term" data-term="antigen" data-term-def="A substance that stimulates the production of an antibody by the immune system. Antigens include toxins, bacteria, foreign blood cells, and&hellip;" data-term-url="/en/glossary/view/antigen/5280">antigen</mark>. If they could do this, it would mean that, by using a common procedure, antibodies could be made to recognize and bind virtually anything chosen as an antigen. The mouse immune system was known to react strongly to sheep red blood <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark> (SRBCs), meaning that mice easily make antibodies against SRBCs. Additionally, antibodies against SRBCs could be detected easily in laboratory tests used routinely in the 1970s. For these reasons, the team chose SRBCs for the test antigen. The goal was to create a lymphocyte-myeloma hybridoma that manufactured antibodies against SRBCs.</p><p>In January 1975, the project succeeded in creating hybridoma <mark class="term" data-term="cell" data-term-def="The basic structural unit of all living things." data-term-url="/en/glossary/view/cell/8286">cells</mark> that not only made antibodies against SRBCs but made them in huge quantities. In their 1975 scientific paper, published in the journal <em>Nature</em>, Milstein and Köhler emphasized the importance of their achievement by discussing potential applications of the hybridoma <mark class="term" data-term="antibody" data-term-def="A Y-shaped protein molecule that is produced by the immune system in response to infection by an antigen. Different antigens provoke&hellip;" data-term-url="/en/glossary/view/antibody/5279">antibody</mark> technology to industry and medicine. Antibodies produced this way are called <em>monoclonal antibodies</em>, because they recognize one <mark class="term" data-term="antigen" data-term-def="A substance that stimulates the production of an antibody by the immune system. Antigens include toxins, bacteria, foreign blood cells, and&hellip;" data-term-url="/en/glossary/view/antigen/5280">antigen</mark> (in fact, they recognize one part of a specific antigen, called an <em>epitope</em>). This is in contrast with <em>polyclonal antibodies</em>, which are made naturally by the immune system when it is stimulated by antigen exposure. In the case of a polyclonal immune response, the antibodies made vary in terms of the physical form and characteristics for binding antigens. In contrast, monoclonal antibodies are all exactly the same and recognize the exact same <mark class="term" data-term="molecule" data-term-def="A particle formed by the chemical bonding of two or more atoms. The molecule is the smallest particle of a&hellip;" data-term-url="/en/glossary/view/molecule/1518">molecule</mark> and the exact same part of a molecule.</p> <div class="comprehension-checkpoint margin-y-4"> <h6 class="comprehension-checkpoint__header"> <span> <span class="icon icon-question"></span> </span> Comprehension Checkpoint </h6> <form class="" name="cc8370"> <div class="form-entry"> <div class="form-entry__field"> <span class="form-entry__field__label">To make antibodies, regular immune cells use genes from</span> <div class="form-entry__option"> <div class="form-entry__option__radio" data-answer="correct"> <label> <input id="q1-8370-0-option-a" name="quiz-option-8370" type="radio" value="only one parent." > <span class="option__label"> <span class="screen-reader-only">a.</span> only one parent. </span> </label> <span class="quiz__response" id="response-8370-0"> <strong>Correct!</strong> </span> </div> <div class="form-entry__option__radio" data-answer="incorrect"> <label> <input id="q1-8370-1-option-b" name="quiz-option-8370" type="radio" value="both parents." > <span class="option__label"> <span class="screen-reader-only">b.</span> both parents. </span> </label> <span class="quiz__response" id="response-8370-1"> <strong>Incorrect.</strong> </span> </div> </div> </div> </div> </form> </div> </section> <section id="toc_6"> <h2>Monoclonal antibodies revolutionize healthcare</h2><p>Because of this specificity of monoclonal antibodies and the large numbers of them that can be made with a hybridoma, they can be used as detectors in medicine and biology. Thus, during the 1980s and 1990s, they revolutionized health care, the pharmaceutical industry, and biology research. Virtually every test for something biological – a blood test for HIV (the <mark class="term" data-term="virus" data-term-def="An ultramicroscopic infectious agent that can only replicate within the cells of living hosts, mainly bacteria, plants, and animals. Viruses have&hellip;" data-term-url="/en/glossary/view/virus/5302">virus</mark> that causes AIDS), hepatitis, influenza, etc. – depends on some kind of monoclonal antibodies, manufactured with the techniques developed by Milstein and his colleagues. Antibodies produced this way also are used for infectious agents in public health settings, detection for defense against biological weapons, and have even been studied for use in detection of life on Mars and other planets. They’re also being employed ever more frequently as therapies against disease, including Ebola.</p> <div class="figure"> <figure> <button class="lightbox-button lightbox-button--icon" data-lightbox="" data-lightbox-src="/img/library/large_images/image_8382.jpg"> <img src="/img/library/modules/mid220/Image/VLObject-8382-150705070754.jpg" alt="Figure 8: César Milstein and Georges Köhler, along with Niels Kaj Jerne (not pictured), won the Nobel Prize in Physiology or Medicine in 1984." /> </button> <figcaption> <p><strong>Figure 8</strong>: César Milstein and Georges Köhler, along with Niels Kaj Jerne (not pictured), won the Nobel Prize in Physiology or Medicine in 1984.</p> <span class="credit">image ©Celia Milstein/MRC Laboratory of Molecular Biology</span> </figcaption> </figure> </div> <p>Though perhaps not realizing just how far into the future Milstein’s work could take medicine, the scientific community understood the importance of the discovery fairly early. Thus, in 1984, together with Köhler and another colleague, Milstein was awarded the <mark class="term" data-term="Nobel Prize" data-term-def="Awards made annually, beginning in 1901, from funds originally established by Alfred B. Nobel for outstanding achievement in physics, chemistry, medicine&hellip;" data-term-url="/en/glossary/view/Nobel+Prize/3843">Nobel Prize</mark> for Physiology and Medicine. </p> </div> </section> <hr class="border-color-dark" /> <footer class="module__footer"> <p class="citation"> <em> David Warmflash, MD “Revolutionizing Medicine with Monoclonal Antibodies” Visionlearning Vol. SCIRE-2 (3), 2015. </em> </p>  <div class="title-list" id="refs" name="refs"> <p class="h6 title-list__title"> References </p> <ul class="title-list__list"> <li><p>American Association of Immunologists. (n.d.). “César Milstein, Ph.D. (1927–2002).” Retrieved May 20, 2015</li> <li><a href="http://www.aai.org/about/History/Notable_Members/Nobel/Milstein_Cesar.html">http://www.aai.org/about/History/Notable_Members/Nobel/Milstein_Cesar.html</a>.</li> <li>Marks, L. (n.d.). “The Story of César Milstein and Monoclonal Antibodies.” Retrieved May 20, 2015</li> <li><a href="http://www.whatisbiotechnology.org/exhibitions/milstein">http://www.whatisbiotechnology.org/exhibitions/milstein</a>.</li> <li>Nobel Media. (2014). “The Nobel Prize in Physiology or Medicine 1984.” Retrieved May 20, 2015</li> <li><a href="http://www.nobelprize.org/nobel_prizes/medicine/laureates/1984/">http://www.nobelprize.org/nobel_prizes/medicine/laureates/1984/</a>.</li> <li>Times Higher Education. (2002). “César Milstein.” Retrieved May 20, 2015</li> <li><a href="http://www.timeshighereducation.co.uk/168036.article">http://www.timeshighereducation.co.uk/168036.article</a>.</p></li> </ul> </div> </footer> </div>   </div>  <div class="order-1 order-2--lg module__tools"> <div class="narrow margin-x-auto position-sticky-top font-size-md"> <div class="padding-2 border-radius box-shadow-1--lg"> <div class="tabs" role="tablist"> <nav> <button class="button button--icon-label" id="tab-button-in-this-module" aria-label="Table of Contents" aria-controls="tab-panel-module__tools" aria-selected="true" role="tab"> <span class="icon icon-list" aria-hidden="true"></span> <span class="button__text">Contents</span> </button> <button class="button button--icon-label" id="tab-button-toggle-terms" aria-controls="tab-panel-toggle-terms" aria-selected="false" role="tab"> <span class="icon icon-glossary-highlight"></span> <span class="button__text">Glossary Terms</span> </button> </nav> <hr class="divider" /> <div class="tabs__panel shown" id="tab-panel-module__tools" aria-labelledby="tab-button-module__tools" role="tabpanel"> <p class="font-weight-bold margin-bottom-1"> Table of Contents </p> <div class="table-of-contents" id="module-toc"> <ul> <li><a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220#toc_1">Snake venom as inspiration</a> </li> <li><a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220#toc_2">A student and activist in Buenos Aires</a> </li> <li><a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220#toc_3">Professional advances, but more political difficulties</a> </li> <li><a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220#toc_4">Antibody research</a> </li> <li> <ul> <li><a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220#toc2_1">Tackling research problems</a> </li> </ul> </li> <li> <ul> <li><a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220#toc2_2">Searching for mutations</a> </li> </ul> </li> <li><a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220#toc_5">A new type of cell: Hybridomas</a> </li> <li><a href="/en/library/scientists-and-research/58/revolutionizing-medicine-with-monoclonal-antibodies/220#toc_6">Monoclonal antibodies revolutionize healthcare</a> </li> </ul> </div> </div>   <div class="tabs__panel" id="tab-panel-toggle-terms" aria-labelledby="tab-button-toggle-terms" role="tabpanel"> <div class="reading-toggle"> <div class="reading-toggle__switch"> <div class="form-entry__option__switch"> <label> <input type="checkbox" name="termsToggleSwitch" id="terms-toggle-switch" /> <span class="switch__slider"></span> <span class="option__label text-decoration-none font-size-md"> Highlight Glossary Terms </span> </label> </div> </div> <div class="reading-toggle__help"> <p> <em> Activate glossary term highlighting to easily identify key terms within the module. 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Revolutionizing Medicine with Monoclonal Antibodies | Scientists and Research | Visionlearning