D A AUTHORITATIVE AND UP TO DATE, this wry and N easy-to-understand book offers an accessible and A GRAPHIC entertaining look at the history, science, and intrigue A G GUIDE behind DNA, the molecule that shook the world. R A P to the H “A UNIQUE, RICHLY DETAILED, and fun biography of DNA grounded IC in deep historical and philosophical knowledge. Israel Rosenfield, G U I Edward Ziff, and Borin Van Loon give us everything we need to D MOLECULE SHOOK WORLD E that the know about biology’s most important molecule.” t o t —OLIVER SACKS, Columbia University, author of The Mind’s Eye and The Man h e Who Mistook His Wife for a Hat M O L E C “THIS IS A REMARKABLE BOOK on DNA, the molecule that lies U L at the core of genetics. The book is novel, easy to read, and fun E t and combines excellent cartoons with good personal vignettes and h a t history. It can be read on its own or as a supplement to a S H conventional textbook. I spent many years mastering genetics O O yet learned new and valuable things from this book. Take a look; K t you will not be disappointed.” h e W —ROBERT TRIVERS, Rutgers University, winner of the Crafoord Prize in O R Biosciences L D Israel Rosenfield, professor at the City University of New York, Guggenheim Fellow, and longtime contributor to the New York Review of Books, has written extensively about evolution, genetics, artificial intelligence, the brain, and memory. Ro se n Edward Ziff, professor of biochemistry and neural science at the New York University fi e School of Medicine, was a collaborator of DNA pioneer Fred Sanger and has researched gene ld structure, transcription, oncogenes, and the brain. , Z i f f Rosenfield and Ziff met while students at Princeton University and have collaborated on essays , & centering on evolution and neuroscience. V a n L Borin Van Loon has been a leading freelance illustrator since 1977. He has designed and o o illustrated fifteen documentary comic books on subjects from Darwin to psychotherapy and n from the Buddha to statistics. He lives and works near London. PRINTED IN THE U.S.A. Cover image: Borin Van Loon Cover design: Milenda Nan Ok Lee C o lu m Columbia University Press / New York b www.cup.columbia.edu ia Israel Rosenfield, Edward Ziff, & Borin Van Loon A GRAPHIC GUIDE to the MolECUlE that SHook the WoRlD ii ii Columbia u niversity Press n ew york A GRAPHIC GUIDE to the MolECUlE that SHook the WoRlD Israel Rosenfield, Edward Ziff, & Borin Van Loon Columbia u niversity Press n ew york Columbia university Press Publishers Since 1893 new york Chichester, west sussex Copyright © 2011 Columbia university Press all rights reserved library of Congress Cataloging-in-Publication Data rosenfield, israel, 1939– ir: to Catherine Dna : a graphic guide to the molecule that shook the world / israel rosenfield, edward Ziff, and borin van loon. p. cm. eZ: to sue, sara, and ben includes bibliographical references. isbn 978-0-231-14270-0 (cloth : alk. paper) bvl: to mandy, simon, Jim, and louisa isbn 978-0-231-14271-7 (pbk. : alk. paper) isbn 978-0-231-51231-2 (e-book) 1. Dna—Popular works. 2. Graphic novels. i. Ziff, edward. ii. van loon, borin, 1951– iii. title. QP624.r67 2011 572.8’6—dc22 2010020889 Columbia university Press books are printed on permanent and durable acid-free paper. this book is printed on paper with recycled content. Printed in the united states of america c 10 9 8 7 6 5 4 3 2 1 p 10 9 8 7 6 5 4 3 2 1 some of this material has appeared in a different form in the New York Review of Books. image on p. ii: Ladder of Ascent / art resource. Columbia university Press Publishers Since 1893 new york Chichester, west sussex Copyright © 2011 Columbia university Press all rights reserved library of Congress Cataloging-in-Publication Data rosenfield, israel, 1939– ir: to Catherine Dna : a graphic guide to the molecule that shook the world / israel rosenfield, edward Ziff, and borin van loon. p. cm. eZ: to sue, sara, and ben includes bibliographical references. isbn 978-0-231-14270-0 (cloth : alk. paper) bvl: to mandy, simon, Jim, and louisa isbn 978-0-231-14271-7 (pbk. : alk. paper) isbn 978-0-231-51231-2 (e-book) 1. Dna—Popular works. 2. Graphic novels. i. Ziff, edward. ii. van loon, borin, 1951– iii. title. QP624.r67 2011 572.8’6—dc22 2010020889 Columbia university Press books are printed on permanent and durable acid-free paper. this book is printed on paper with recycled content. Printed in the united states of america c 10 9 8 7 6 5 4 3 2 1 p 10 9 8 7 6 5 4 3 2 1 some of this material has appeared in a different form in the New York Review of Books. image on p. ii: Ladder of Ascent / art resource. Contents Contents Preface xi Introduction 1 A Brief History of Genetics 5 What Is DNA? 28 The New Biology 34 DNA Replication 58 What Information Is Stored in a Gene? 60 Where Did the Notion of Messenger RNA Come From? 67 Transcription 70 Translation 75 Genetic Code 85 PaJaMo and the Operon 88 The Diversity of Gene Expression 95 A Brief Reminder of Relative Size 102 A Brief Reminder of Genes in Humans 104 Prokaryotes versus Eukaryotes 105 Restriction Enzymes and Genetic Engineering 114 Cloning and Sequencing Genes 124 Exons, Introns, and Splicing 136 Chromatin and Histones 143 Gene Families 144 Controlling Genes for Antibodies 146 Chromosomes 154 Telomeres 155 Imprinting and Micro RNAs: “Hidden” Layers of Gene Regulation 156 Epigenetics 157 Prions 159 The Human Genome Project 161 The Human Genome Unveiled 163 High Throughput Sequencing 165