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. ' W " . ^ BIOL ^H^^'i<i

THIRD EOmON

ICAL SCIENCE SCOTT FREEMAN

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BIOLOGICAL SCIENCE, 3e

Study Key

O Key Concept Important Information

• PracticeIt •QUESTION •EXERCISE

cnecJCfouMuraerstarTom^ ^ ^ ^ ^ ^ H 1 Ifyou understand that... O 1

1 You should be able to... 1 •

1

The Gold Thread helps you pick out important information.

The Blue Thread helps you practice what you've learned.

Check Your Understanding boxes review important information and ask you to apply it.

Pointing Hand — ^ ' ^

SUMMARY TABLE

BIOSKILLS

"Pointers" help you focus on important parts of figures.

Summary Tables help you review important information. Complete list on inside back cover.

Experiment Boxes help you learn how scientists work with data.

• BioSkills teach you fundamental skills used by scientists. Complete list on inside back cover.

wwwjnasteringbio.coni is also your resource for • Answers to text, table, and figure caption questions and exercises • Answers to Check Your Understanding boxes • Online study guides and quizzes • Additional study tools including the E-Book for Biological Science 3rd ed., textbook art, animations, and videos.

(MB) EVraI¥:TifTTiffT[«iil at www.masteringbio.com

(MB) BioFtuc atwww.masteringbio.com

Online Resources extend the textbook's learning tools.

Web Animations help you review important textbook topics.

BioFlix 3D Animations help you visualize difficult biology topics.

M P I C l " P r T n n D I r\t%\ r* "M ^ ° "^ gateway to the most widely-used online I l a D L e i m y t J I U L ^ i J T biology self-study materials and more...

With your purchase of a new copy of Freeman's Biological Science, Third Edition, you should have received a Student Access Kit for MasteringBiology^**. This kit contains instructions and a code for accessing this dynamic website. Your Student Access Kit looks like this:

D O N ' T T H R O W YOUR ACCESS KIT AWAY!

Ifyou did not purchase a new textbook or cannot locate the Student Access Kit, you can purchase your subscription to MasteringBiology^*^ online with a major credit card by selecting Buy Now. To do so, go to www.masteringbio.com. Identify the exact title and edition of your textbook when prompted.

I N S T R U C T O R S :

Look for your "Instructor Access Kit" that you received with your bool<. Contact

your Pearson Benjamin Cummings Sales Representative ifyou need assistance.

MasteringBiOLOGY What is MasteringBiology™?

MasteringBiology''''^ offers two valuable learning systems:

1. The Study Area for studying on your own or in a study group

2. MasteringBiology^** Assignments that your instructor may require

REQUIRED for access to online materials

STUDENT ACCESS KIT Register online at www.masteringbio.com

Mastering^ BIOLOGY

''Sl0^^^^- THIRD EDITION

BIOLOGICAL SCIENCE SCOTT FREEMAN

The Study Area provides state-of-the-art BioFlixT"^ 3-D Animations, Discovery videos, Web Animations, Cell Biology videos, interactive eBook, answers to textbook questions and much more (see other side).

MasteringBiology'''** Assignments may include Tutorials and other Multiple Choice Questions required by your instructor. MasteringBiology^^ tutorials coach you with feedback specific to your needs and offer hints when you get stuck, optimizing your study time and maximizing your learning.

@ Features of the MasteringBiology^^ Study Area for Freeman's Biological Science, Third Edition

C H A P T E R G U I D E Prepare for the test by reviewing important information outlined in the Chapter Guide. The Chapter Guide provides a quick overview of key concepts, practice quizzes, and review questions to help students study for the exam. Link directly to the BioFlix^^ 3-D animations. Discovery Videos, BioSkills, Cell Biology Animations, and interactive eBook to help students master biology's most difficult topics.

BioFlix

B I O F L I X ™ 3 - D A N I M A T I O N S

BioFhx''^'^ are 3-D movie-quahty animations with carefiiUy con­

structed student tutorials, labeled shde shows, study sheets, and quizzes which help bring biology to hfe. Topics include tour of an animal cell, tour of a plant cell, cellular respiration, photosynthesis, mito­sis, meiosis, protein synthesis, water transport in plants, how neurons work, and muscle contraction.

V I D E O S Watch brief Discovery ChanneF"^ video clips on 29 different topics including antibiotic resistance,

fighting cancer, and introduced species.

DLi:coueru

W E B A N I M A T I O N S WI33M113MM Learn about biological concepts in a visual format with online web animations. Animations include pre-quizzes and post-quizzes to test students' understanding of dynamic concepts and processes. Additional Cell Biology Animations provide vivid images of the fiinctions and processes of the cell.

G R A P H I T ! Graphing tutorials show students how to plot, interpret, and critically evaluate real data.

C H A P T E R Q U I Z Z E S Students can assess their comprehension of material with 20 multiple-choice quiz questions for each chapter.

A N S W E R K E Y Find answers to the book's figure and table cap­tion questions and exercises, "you should be able to . . . " activities, and end-of-chapter questions.

A R T View and print artwork from the textbook to foUow along in class or use as a study tool in review sessions.

A U D I O G L O S S A R Y Build your biology vocabulary with our audio glossary. Hear the correct pronunciation and learn the meaning of all key terms.

E - B O O K

Refer to a convenient ordine version of the book while you study.

BIOLOGICAL SCIENCE

k

Actual Size: 4 cm •J

ABOUT THE PHOTO: He'e, Octopus sp. (juvenile), photographed at Midway Atoll National Wildlife Refuge, located in the Northwestern Hawaiian Islands, 29 IVIarch 2003, by David Liittschwager and Susan Middleton, authors of Archipelago: Portraits of Life in tfie World's Most Remote Island Sanctuary (National Geographic Society, 2005).

BIOLOGICAL SCIENCE THIRD EDITION

SCOTT FREEMAN University of Washington

PEARSON

Benjamin Cummings

San Francisco Boston New York Cape Town Hong Kong London Madrid Mexico City Montreal Munich Paris Singapore Sydney Tokyo Toronto

Editor-in-Chief, Biology: Beth Wilbur Sponsoring Editors: Andrew Gilfiilan, Susan Winslow,

Becky Ruden Editorial Project Managers; Ann Heath, Sonia DiVittorio Development Editors: Erin Mulligan,

Mary Catherine Hager, Susan Weisberg Senior Production Supervisor: Shannon Tozier Associate Editor: Mercedes Grandin Assistant Editor: Anna Amato Editorial Assistants: Lisa Tarabokjia, Nina Sparer Media Producer: Ericka G'Benar Executive Directors of Development: Carol Trueheart,

Deborah Gale Executive Marketing Manager: Lauren Harp Director of Marketing; Christy Lawrence Managing Editor: Mike Early

Executive Managing Editor: Erin Gregg Production Service and Composition: Pre-Press PMG Illustrations: Kim Quillin, Imagineering Media Services, Inc.,

Greg Williams Interior Design: Marilyn Perry Cover Design: Kim Quillin, Marilyn Perry Cover Production: Laura Wieglab and Side by Side Studios Manufacturing Buyer: Mike Early and Michael Pene Director, Image Resource Center: Melinda Patelli Manager, Rights and Permissions: Zina Arabia Visual Research Coordinator: Elaine Soares Image Permission Coordinator: Debbie Latronica Photo Researcher: Yvonne Gerin Cover Printer: Phoenix Color Printer and Binder: R.R. Donnelley, Willard

Cover Image: Octopus-he'e, Octopus sp. (juvenile); David Liittschwager/Susan Middleton

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings, 1301 Sansome St., San Francisco, CA 94111. All rights reserved. Manufactured in the United States of America. This publication is protected by Copyright and permission should be obtained from the publisher prior to any prohibited reproduction, storage in a retrieval system, or transmission in any form or by any means, electronic, mechanical, photocopying, recording, or likewise. To obtain permis-sion(s) to use material from this work, please submit a written request to Pearson Education, Inc., Permissions Department, 1900 E. Lake Ave., Glenview, IL 60025. For information regarding permissions, call (847) 486-2635.

Many of the designations used by manufacturers and sellers to distinguish their products are claimed as trademarks. Where those designations appear in this book, and the publisher was aware of a trademark claim, the designations have been printed in initial caps or all caps.

MasteringBiology™ and BioFlix™ are trademarks, in the U.S. and/or other countries, of Pearson Education, Inc. or its affiliates.

Library of Congress Cataloging-in-Publication Data

Freeman, Scott Biological science / Scott Freeman.—3rd ed.

p. cm. Includes index. ISBN 0-13-224950-2 (student edition)—ISBN 0-13-224985-5 (professional

copy)—ISBN 0-13-225308-9 (volume 1)—ISBN 0-13-232543-8 (volume 2) —ISBN 0-13-156816-7 (volume 3)

1. Biology. I. Title. QH308. 2. F73 2007 570—dc22

2007036866

ISBN 10-digit 0-13-224950-2; 13-digit 978-0-13-224950-8 (Student edition) ISBN 10-digit 0-13-224985-5; 13-digit 978-0-13-224985-0 (Professional copy) ISBN 10-digit 0-13-225308-9; 13-digit 978-0-13-225308-6 (Volume 1) ISBN 10-digit 0-13-232543-8; 13-digit 978-0-13-232543-1 (Volume 2) ISBN 10-digit 0-13-156816-7; 13-digit 978-0-13-156816-7 (Volume 3)

PEARSON

Ben.jai.iin -Cummings 2 3 4 5 6 7 8 9 10—DOW—11 10 09 08

www.pearsonhighered.com

Brief Contents

1 Biology and the Tree of Life 1

THE MOLECULES OF LIFE 18

2 Water and Carbon:The Chemical Basis of Life 18

3 Protein Structure and Function 43

4 Nucleic Acids and the RNA World 67

5 An Introduction to Carbohydrates 82

6 Lipids, Membranes, and the First Cells 95

CELL STRUCTURE AND FUNCTION 119

7 Inside the Cell 119

8 Cell-Cell Interactions 150

9 Cellular Respiration and Fermentation 170

10 Photosynthesis 198

11 The Cell Cycle 222

GENE STRUCTURE AND EXPRESSION 243

12 Meiosis 243

13 Mendel and the Gene 265

14 DNAand the Gene: Synthesis and Repair 295

15 How Genes Work 316

16 Transcription and Translation 329

17 Control of Gene Expression in Bacteria 352

18 Controlof Gene Expression in Eukaryotes 370

19 Analyzing and Engineering Genes 389

20 Genomics 415

DEVELOPMENTAL BIOLOGY 434

21 Principles of Development 434

22 An Introduction to Animal Development 451

23 An Introduction to Plant Development 467

EVOLUTIONARY PROCESSES AND PATTERNS 481

24 Evolution by Natural Selection 481

25 Evolutionary Processes 503

26 Speciation 526

27 Phylogenies and the History of Life 543

THE DIVERSIFICATION OF LIFE 566

28 Bacteria and Archaea 566

29 Protists 593

30 Green Plants 626

31 Fungi 664

32 An Introduction to Animals 688

33 Protostome Animals 712

34 Deuterostome Animals 737

35 Viruses 769

HOW PLANTS WORK 791

36 Plant Form and Function 791

37 Water and Sugar Transport in Plants 813

38 Plant Nutrition 837

39 Plant Sensory Systems, Signals, and Responses 857

40 Plant Reproduction 892

HOW ANIMALS WORK 913

41 Animal Form and Function 913

42 Water and Electrolyte Balance in Animals 934

43 Animal Nutrition 957

44 Gas Exchange and Circulation 978

45 Electrical Signals in Animals 1006

46 Animal Sensory Systems and Movement 1030

47 Chemical Signals in Animals 1054

48 Animal Reproduction 1079

49 The Immune System in Animals 1104

ECOLOGY 1125

50 An Introduction to Ecology 1125

51 Behavior 1149

52 Population Ecology 1173

53 Community Ecology 1196

54 Ecosystems 1222

55 Biodiversity and Conservation Biology 1244