iii

iv

PUBLISHER: Katherine Ahr Parker

ACQUISITIONS EDITOR: Beth Cole

DEVELOPMENTAL EDITORS: Erica Champion, Heather Moffat

EDITORIAL ASSISTANTS: Nandini Ahuja, Abigail Fagan

EXECUTIVE MARKETING MANAGER: Will Moore

SENIOR PROJECT EDITOR: Elizabeth Geller

DESIGN MANAGER: Blake Logan

TEXT DESIGNER: Patrice Sheridan

COVER DESIGN: Blake Logan

ILLUSTRATION COORDINATOR: Janice Donnola

ART DEVELOPMENT EDITOR: H. Adam Steinberg, Art for Science

PERMISSIONS MANAGER: Jennifer MacMillan

PHOTO EDITOR: Sheena Goldstein

PHOTO RESEARCHER: Teri Stratford

TEXT PERMISSIONS: Felicia Ruocco, Hilary Newman

MEDIA AND SUPPLEMENTS EDITORS: Amy Thorne, Kathleen Wisneski

SENIOR MEDIA PRODUCER: Chris Efstratiou

SENIOR PRODUCTION SUPERVISOR: Paul Rohloff

COMPOSITION: codeMantra

PRINTING AND BINDING: RR Donnelley

COVER IMAGE: Dr. Tomas Kirchhausen and Dr. Lei Lu

ABOUT THE COVER: Imaging of the intracellular organelles of a live human HeLa cell shows the dramatic morphological changes that accompany the process of cell division. The membrane of the endoplasmic reticulum (ER) is labeled green by a fluorescently tagged component of the translocon (GFP-Sec61β) and chromatin is labeled red by a fluorescently tagged histone (H2B-mRFP). Front: An interphase cell showing uncondensed chromatin filling the nucleus, with the ER as a reticulum of cisternae surrounding the nucleus and interconnected with lace-like tubules at the cell periphery. Back: Prior to cell division the chromatin condenses to reveal the worm-like structure of individual chromosomes, the nuclear envelope breaks down, and the ER condenses into an array of cisternae surrounding the condensed chromosomes. As cell division proceeds the replicated chromosomes will segregate equally into two daughter cells, nuclear envelopes will form in the daughter cells, and the ER will return to its characteristic reticular organization. Cover photo: Dr. Tomas Kirchhausen & Dr. Lei Lu.

Library of Congress Control Number: 2015957295

ISBN-13: 978-1-4641-8339-3

ISBN-10: 1-4641-8339-2

© 2016, 2013, 2008, 2004 by W. H. Freeman and Company

All rights reserved.

Printed in the United States of America

First printing

W. H. Freeman and Company

One New York Plaza, Suite 4500, New York, NY 10004-1562

www.macmillanhighered.com

v

TO OUR STUDENTS AND TO OUR TEACHERS,

from whom we continue to learn,

AND TO OUR FAMILIES,

for their support, encouragement, and love