From Genes to Genomes
HARTWELL ~ GOLDBERG ~ FISCHER ~ HOOD
, SIXTH E D I TI O N
Genetics
From Genes to Genomes
Leland H. Hartwell
FRED HUTCHISON CANCER CENTER
Michael L. Goldberg
CORNELL UNIVERSITY
Janice A. Fischer
UNIVERSITY OF TEXAS AT AUSTIN
Leroy Hood
THE INSTITUTE FOR SYSTEMS BIOLOGY
,GENETICS: FROM GENES TO GENOMES, SIXTH EDITION
Published by McGraw-Hill Education, 2 Penn Plaza, New York, NY 10121. Copyright © 2018 by
McGraw-Hill Education. All rights reserved. Printed in the United States of America. Previous editions
© 2015, 2011, and 2008. No part of this publication may be reproduced or distributed in any form or by
any means, or stored in a database or retrieval system, without the prior written consent of McGraw-Hill
Education, including, but not limited to, in any network or other electronic storage or transmission, or
broadcast for distance learning.
Some ancillaries, including electronic and print components, may not be available to customers outside the
United States.
This book is printed on acid-free paper.
1 2 3 4 5 6 7 8 9 LWI 21 20 19 18 17
ISBN 978–1–259–70090–3
MHID 1–259–70090–9
Portfolio Manager: Justin K.Wyatt, PhD
Lead Product Developer: Fran Simon
Senior Marketing Manager: Kelly Brown
Senior Content Project Manager: Vicki Krug
Lead Content Project Manager: Christina Nelson
Buyer: Jennifer Pickel
Lead Designer: David Hash
Lead Content Licensing Specialist: Lori Hancock
Content Licensing Specialist: Lori Slattery
Cover Image: © Sydney F. Vize
Design Element: Front and Back Matter DNA helix header, © MedicalRF.com
Compositor: Aptara, Inc
Printer: LSC Communications
All credits appearing on page or at the end of the book are considered to be an extension of the copyright page.
Library of Congress Cataloging-in-Publication Data
Names: Hartwell, Leland, author.
Title: Genetics : from genes to genomes / Leland Hartwell, Michael Goldberg,
Janice Fischer, Lee Hood, Charles F. Aquadro, Bruce Bejcek.
Description: Sixth edition. ∣ New York, NY : McGraw-Hill Education, [2018] ∣
Includes index.
Identifiers: LCCN 2017017175∣ ISBN 9781259700903 (alk. paper) ∣
ISBN 1259700909 (alk. paper)
Subjects: LCSH: Genetics.
Classification: LCC QH430 .G458 2018 ∣ DDC 576.5—dc23 LC record available at
https://lccn.loc.gov/2017017175
The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website
does not indicate an endorsement by the authors or McGraw-Hill Education, and McGraw-Hill Education
does not guarantee the accuracy of the information presented at these sites.
mheducation.com/highered
, About the Authors
Dr. Leland Hartwell is President and Director of Seattle’s Fred Hutchinson Cancer Research
Center and Professor of Genome Sciences at the University of Washington.
Dr. Hartwell’s primary research contributions were in identifying genes that control cell divi-
sion in yeast, including those necessary for the division process as well as those necessary for the
fidelity of genome reproduction. Subsequently, many of these same genes have been found to
control cell division in humans and often to be the site of alteration in cancer cells.
Dr. Hartwell is a member of the National Academy of Sciences and has received the Albert
Lasker Basic Medical Research Award, the Gairdner Foundation International Award, the Genetics
Society Medal, and the 2001 Nobel Prize in Physiology or Medicine.
Dr. Michael Goldberg is a Professor at Cornell University, where he teaches introductory genet-
ics and human genetics. He was an undergraduate at Yale University and received his Ph.D. in
biochemistry from Stanford University. Dr. Goldberg performed postdoctoral research at the
Biozentrum of the University of Basel (Switzerland) and at Harvard University, and he received
an NIH Fogarty Senior International Fellowship for study at Imperial College (England) and fel-
lowships from the Fondazione Cenci Bolognetti for sabbatical work at the University of Rome
(Italy). His current research uses the tools of Drosophila genetics and the biochemical analysis of
frog egg cell extracts to investigate the mechanisms that ensure proper cell cycle progression and
chromosome segregation during mitosis and meiosis.
Dr. Janice Fischer is a Professor at The University of Texas at Austin, where she is an award-
winning teacher of genetics and Director of the Biology Instructional Office. She received her
Ph.D. in biochemistry and molecular biology from Harvard University, and did postdoctoral
research at The University of California at Berkeley and The Whitehead Institute at MIT. In her
research, Dr. Fischer used Drosophila first to determine how tissue-specific transcription works,
and then to examine the roles of ubiquitin and endocytosis in cell signaling during development.
Dr. Lee Hood received an M.D. from the Johns Hopkins Medical School and a Ph.D. in bio-
chemistry from the California Institute of Technology. His research interests include immunol-
ogy, cancer biology, development, and the development of biological instrumentation (for
example, the protein sequencer and the automated fluorescent DNA sequencer). His early research
played a key role in unraveling the mysteries of antibody diversity. More recently he has pioneered
systems approaches to biology and medicine.
Dr. Hood has taught molecular evolution, immunology, molecular biology, genomics and bio-
chemistry and has co-authored textbooks in biochemistry, molecular biology, and immunology, as
well as The Code of Codes—a monograph about the Human Genome Project. He was one of the
first advocates for the Human Genome Project and directed one of the federal genome centers that
sequenced the human genome. Dr. Hood is currently the president (and co-founder) of the cross-
disciplinary Institute for Systems Biology in Seattle, Washington.
Dr. Hood has received a variety of awards, including the Albert Lasker Award for Medical
Research (1987), the Distinguished Service Award from the National Association of Teachers
(1998) and the Lemelson/MIT Award for Invention (2003). He is the 2002 recipient of the Kyoto
Prize in Advanced Biotechnology—an award recognizing his pioneering work in developing the
protein and DNA synthesizers and sequencers that provide the technical foundation of modern
biology. He is deeply involved in K–12 science education. His hobbies include running, mountain
climbing, and reading.
iii
HARTWELL ~ GOLDBERG ~ FISCHER ~ HOOD
, SIXTH E D I TI O N
Genetics
From Genes to Genomes
Leland H. Hartwell
FRED HUTCHISON CANCER CENTER
Michael L. Goldberg
CORNELL UNIVERSITY
Janice A. Fischer
UNIVERSITY OF TEXAS AT AUSTIN
Leroy Hood
THE INSTITUTE FOR SYSTEMS BIOLOGY
,GENETICS: FROM GENES TO GENOMES, SIXTH EDITION
Published by McGraw-Hill Education, 2 Penn Plaza, New York, NY 10121. Copyright © 2018 by
McGraw-Hill Education. All rights reserved. Printed in the United States of America. Previous editions
© 2015, 2011, and 2008. No part of this publication may be reproduced or distributed in any form or by
any means, or stored in a database or retrieval system, without the prior written consent of McGraw-Hill
Education, including, but not limited to, in any network or other electronic storage or transmission, or
broadcast for distance learning.
Some ancillaries, including electronic and print components, may not be available to customers outside the
United States.
This book is printed on acid-free paper.
1 2 3 4 5 6 7 8 9 LWI 21 20 19 18 17
ISBN 978–1–259–70090–3
MHID 1–259–70090–9
Portfolio Manager: Justin K.Wyatt, PhD
Lead Product Developer: Fran Simon
Senior Marketing Manager: Kelly Brown
Senior Content Project Manager: Vicki Krug
Lead Content Project Manager: Christina Nelson
Buyer: Jennifer Pickel
Lead Designer: David Hash
Lead Content Licensing Specialist: Lori Hancock
Content Licensing Specialist: Lori Slattery
Cover Image: © Sydney F. Vize
Design Element: Front and Back Matter DNA helix header, © MedicalRF.com
Compositor: Aptara, Inc
Printer: LSC Communications
All credits appearing on page or at the end of the book are considered to be an extension of the copyright page.
Library of Congress Cataloging-in-Publication Data
Names: Hartwell, Leland, author.
Title: Genetics : from genes to genomes / Leland Hartwell, Michael Goldberg,
Janice Fischer, Lee Hood, Charles F. Aquadro, Bruce Bejcek.
Description: Sixth edition. ∣ New York, NY : McGraw-Hill Education, [2018] ∣
Includes index.
Identifiers: LCCN 2017017175∣ ISBN 9781259700903 (alk. paper) ∣
ISBN 1259700909 (alk. paper)
Subjects: LCSH: Genetics.
Classification: LCC QH430 .G458 2018 ∣ DDC 576.5—dc23 LC record available at
https://lccn.loc.gov/2017017175
The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website
does not indicate an endorsement by the authors or McGraw-Hill Education, and McGraw-Hill Education
does not guarantee the accuracy of the information presented at these sites.
mheducation.com/highered
, About the Authors
Dr. Leland Hartwell is President and Director of Seattle’s Fred Hutchinson Cancer Research
Center and Professor of Genome Sciences at the University of Washington.
Dr. Hartwell’s primary research contributions were in identifying genes that control cell divi-
sion in yeast, including those necessary for the division process as well as those necessary for the
fidelity of genome reproduction. Subsequently, many of these same genes have been found to
control cell division in humans and often to be the site of alteration in cancer cells.
Dr. Hartwell is a member of the National Academy of Sciences and has received the Albert
Lasker Basic Medical Research Award, the Gairdner Foundation International Award, the Genetics
Society Medal, and the 2001 Nobel Prize in Physiology or Medicine.
Dr. Michael Goldberg is a Professor at Cornell University, where he teaches introductory genet-
ics and human genetics. He was an undergraduate at Yale University and received his Ph.D. in
biochemistry from Stanford University. Dr. Goldberg performed postdoctoral research at the
Biozentrum of the University of Basel (Switzerland) and at Harvard University, and he received
an NIH Fogarty Senior International Fellowship for study at Imperial College (England) and fel-
lowships from the Fondazione Cenci Bolognetti for sabbatical work at the University of Rome
(Italy). His current research uses the tools of Drosophila genetics and the biochemical analysis of
frog egg cell extracts to investigate the mechanisms that ensure proper cell cycle progression and
chromosome segregation during mitosis and meiosis.
Dr. Janice Fischer is a Professor at The University of Texas at Austin, where she is an award-
winning teacher of genetics and Director of the Biology Instructional Office. She received her
Ph.D. in biochemistry and molecular biology from Harvard University, and did postdoctoral
research at The University of California at Berkeley and The Whitehead Institute at MIT. In her
research, Dr. Fischer used Drosophila first to determine how tissue-specific transcription works,
and then to examine the roles of ubiquitin and endocytosis in cell signaling during development.
Dr. Lee Hood received an M.D. from the Johns Hopkins Medical School and a Ph.D. in bio-
chemistry from the California Institute of Technology. His research interests include immunol-
ogy, cancer biology, development, and the development of biological instrumentation (for
example, the protein sequencer and the automated fluorescent DNA sequencer). His early research
played a key role in unraveling the mysteries of antibody diversity. More recently he has pioneered
systems approaches to biology and medicine.
Dr. Hood has taught molecular evolution, immunology, molecular biology, genomics and bio-
chemistry and has co-authored textbooks in biochemistry, molecular biology, and immunology, as
well as The Code of Codes—a monograph about the Human Genome Project. He was one of the
first advocates for the Human Genome Project and directed one of the federal genome centers that
sequenced the human genome. Dr. Hood is currently the president (and co-founder) of the cross-
disciplinary Institute for Systems Biology in Seattle, Washington.
Dr. Hood has received a variety of awards, including the Albert Lasker Award for Medical
Research (1987), the Distinguished Service Award from the National Association of Teachers
(1998) and the Lemelson/MIT Award for Invention (2003). He is the 2002 recipient of the Kyoto
Prize in Advanced Biotechnology—an award recognizing his pioneering work in developing the
protein and DNA synthesizers and sequencers that provide the technical foundation of modern
biology. He is deeply involved in K–12 science education. His hobbies include running, mountain
climbing, and reading.
iii
