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Contents
1 Introduction 1
Scope and Outlook
1.3 Behavioral Neuroscientists Use
Several Levels of Analysis 10
1.4 The History of Research on the Brain
Machine or Human? 1 and Behavior Begins in Antiquity 14
1.1 The Brain Is Full of Surprises 2 BOX 1.2 Bigger Better? The Case of the Brain
BOX 1.1 We Are All Alike, and We Are and Intelligence 18
All Different 6 The Cutting Edge ■ Behavioral Neuroscience
Is Advancing at a Tremendous Rate 20
1.2 Three Approaches Relate Brain
and Behavior 7 Visual Summary 22

PART I Biological Foundations of Behavior 23

2 Functional Neuroanatomy 25
The Cells and Structures of the
Nervous System
3 Neurophysiology 63
The Generation, Transmission, and
Integration of Neural Signals
A Stimulating Experience 25 The Laughing Brain 63
2.1 S
 pecialized Cells Make Up the 3.1 E
 lectrical Signals Are the Vocabulary
Nervous System 26 of the Nervous System 64
BOX 2.1 Visualizing the Cells of the Brain 32 BOX 3.1 Voltage Clamping and Patch
Clamping 72
2.2 T
 he Nervous System Consists of
Central and Peripheral Divisions 36 BOX 3.2 Changing the Channel 76
BOX 2.2 Three Customary Orientations for 3.2 S
 ynapses Transmit Information from
Viewing the Brain and Body 43 One Neuron to Another 77
2.3 T
 he Brain Shows Regional 3.3 A
 ction Potentials Cause the Release
Specialization of Functions 46 of Transmitter Molecules into the
Synaptic Cleft 81
2.4 S
 pecialized Support Systems Protect
and Nourish the Brain 50 3.4 G
 ross Electrical Activity of the Brain
Is Readily Detected 90
2.5 B
 rain-Imaging Techniques Reveal the
Structure and Function of the Living The Cutting Edge ■ Optogenetics: Using Light
Human Brain 54 to Probe Brain-Behavior Relationships 93
BOX 2.3 Isolating Specific Brain Activity 57 Visual Summary 95
The Cutting Edge ■ Two Heads Are Better
Than One 59
Vi su a l S umm a r y 61
4 The Chemistry of Behavior 97 4.8 S
 ubstance Abuse and Addiction Are
Neurotransmitters and Worldwide Social Problems 127
Neuropharmacology The Cutting Edge ■ Uncovering the Insula 131
The Birth of a Pharmaceutical Problem Visual Summary 135
Child 97
4.1 Synaptic Transmission Involves a
Complex Electrochemical Process 98 5 
Hormones and the Brain 137

4.2 M
 any Neurotransmitters Have Been Crafting a Personality through
Identified 100 Hormones 137
4.3 N
 eurotransmitter Systems Form a 5.1 H
 ormones Have Many Actions
Complex Array in the Brain 101 in the Body 138
BOX 4.1 Pathways for Neurotransmitter 5.2 H
 ormones Have a Variety of Cellular
Synthesis 103 Actions 143
4.4 T
 he Effects of a Drug Depend on Its BOX 5.1 Techniques of Modern Behavioral
Site of Action and Dose 106 Endocrinology 146

4.5 D
 rugs Affect Each Stage of 5.3 E
 ach Endocrine Gland Secretes
Neural Conduction and Synaptic Specific Hormones 149
Transmission 112 The Cutting Edge ■ Bones Secrete
4.6 S
 ome Neuroactive Drugs Ease the Hormones to Regulate Appetite 158
Symptoms of Injury or Psychiatric 5.4 H
 ormones Regulate Social Behaviors
Illness 116 and Vice Versa 158
4.7 S
 ome Neuroactive Drugs Are Used to Visual Summary 164
Alter Conscious Experiences 119

PART II Evolution and Development of


the Nervous System
165

6  volution of the Brain and


E 6.4 T
 he Evolution of Vertebrate Brains
Behavior 167 Reflects Changes in Behavior 181
6.5 M
 any Factors Led to the Rapid
We Are Not So Different, Are We? 167 Evolution of a Large Cortex in
6.1 H
 ow Did the Enormous Variety of Primates 186
Species Arise on Earth? 168 BOX 6.3 Evolutionary Psychology 190
6.2 W
 hy Should We Study Other 6.6 E
 volution Continues Today 193
Species? 174
The Cutting Edge ■ Are Humans Still
BOX 6.1 Why Should We Study Particular Evolving? 194
Species? 175
Visual Summary 197
BOX 6.2 To Each Its Own Sensory World 177
6.3 A
 ll Vertebrate Brains Share the Same
Basic Structures 178

VIII CO NT E NTS
7  ife-Span Development of the
L 7.3 L
 ifelong Synapse Rearrangement
Brain and Behavior 199 Is Guided by Experience 215

Overcoming Blindness 199 7.4 E


 xperience Can Alter Gene
Expression to Affect Brain
7.1 Neural Development Is Guided Development 222
by the Interaction of Genetic and The Cutting Edge ■ Harnessing Glia to
Environmental Factors 200 Reverse an Inherited Brain Disorder 224
7.2 D
 evelopment of the Nervous System 7.5 T
 he Brain Continues to Change
Can Be Divided into Six Distinct as We Grow Older 226
Stages 204
Visual Summary 23 0
BOX 7.1 Transgenic and Knockout Mice 209
BOX 7.2 Degeneration and Regeneration of
Nervous Tissue 211

PART III Perception and Action 231

8 
General Principles of Sensory
Processing, Touch, and Pain
233
9  earing, Balance, Taste, and
H
Smell 269
No Ear for Music 269
What’s Hot? What’s Not? 233
HEARING 270
SENSORY PROCESSING 234
9.1 Pressure Waves in the Air Are
8.1 S
 ensory Receptor Organs Detect Perceived as Sound 270
Energy or Substances 234 BOX 9.1 The Basics of Sound 270
8.2 S
 ensory Information Processing Is 9.2 A
 uditory Signals Run from Cochlea
Selective and Analytical 238 to Cortex 277
BOX 8.1 Synesthesia 245
9.3 P
 itch Information Is Encoded in Two
TOUCH: MANY SENSATIONS BLENDED Complementary Ways 279
TOGETHER 246 9.4 B
 rainstem Auditory Systems
8.3 S
 kin Is a Complex Organ That Are Specialized for Localizing
Contains a Variety of Sensory Sounds 281
Receptors 246 9.5 T
 he Auditory Cortex Processes
PAIN: AN UNPLEASANT BUT ADAPTIVE Complex Sounds 283
EXPERIENCE 252 9.6 H
 earing Loss Is a Major Disorder of
8.4 H
 uman Pain Can Be Measured 252 the Nervous System 286
The Cutting Edge ■ Evolving an Indifference VESTIBULAR PERCEPTION 290
to Toxins 256
9.7 A
 n Inner Ear System Senses Gravity
8.5 P
 ain Can Be Difficult to Control 261 and Acceleration 290
Vi su a l S um ma r y 266 THE CHEMICAL SENSES: TASTE
AND SMELL 293

CONTENTS IX
9.8 C
 hemicals in Foods Are Perceived 10.7 Visual Neuroscience Can Be
as Tastes 293 Applied to Alleviate Some Visual
The Cutting Edge ■ More Than a Matter Deficiencies 340
of Taste 298 The Cutting Edge ■ Seeing the Light 342
9.9 C
 hemicals in the Air Elicit Odor Visual Summary 344
Sensations 299
Visu a l S um m a r y 30 6
11  otor Control and
M
Plasticity 347

10 Vision 309
From Eye to Brain
What You See Is What You Get
11.1 The Behavioral View Considers
347

When Seeing Isn’t Seeing 309 Reflexes versus Plans 348


10.1 The Retina Transduces Light into 11.2 N
 euroscience Reveals Hierarchical
Neuronal Activity 310 Systems 350
10.2 Properties
 of the Retina Shape 11.3 T
 he Spinal Cord Is a Crucial Link in
Many Aspects of Our Vision 314 Controlling Body Movement 357
10.3 N
 eural Signals Travel from the 11.4 P
 athways from the Brain
Retina to Several Brain Regions 318 Control Different Aspects of
Movements 361
BOX 10.1 Eyes with Lenses Have Evolved in
Several Phyla 321 BOX 11.1 Cortical Neurons Can Guide a
Robotic Arm 365
10.4 N
 eurons at Different Levels of the
Visual System Have Very Different 11.5 E
 xtrapyramidal Systems Also
Receptive Fields 322 Modulate Motor Commands 369
10.5 C
 olor Vision Depends on Special The Cutting Edge ■ Cerebellar Glia Play a
Channels from the Retinal Cones Role in Fine Motor Coordination 372
through Cortical Area V4 332 11.6 B
 rain Disorders Can Disrupt
BOX 10.2 Most Mammalian Species Have Movement 373
Some Color Vision 334 BOX 11.2 Prion-Like Neurodegeneration May
10.6 T
 he Many Cortical Visual Areas Be at Work in Parkinson’s 375
Are Organized into Two Major Visual Summary 378
Streams 337

PART IV Regulation and Behavior 381

12 Sex 383
Evolutionary, Hormonal, and Neural
Bases
12.1 R
 eproductive Behavior Can Be
Divided into Four Stages 384

Genitals and Gender: What Makes Us 12.2 T


 he Neural Circuitry of the
Male and Female? 383 Brain Regulates Reproductive
Behavior 388
SEXUAL BEHAVIOR 384

X CO NT E NTS
The Cutting Edge ■ Sexual Experience 13.4 N
 utrient Regulation Helps Prepare
Solidifies Neural Circuits for Mating 390 for Future Needs 429
12.3 T
 he Hallmark of Human Sexual 13.5 A
 Hypothalamic Appetite Controller
Behavior Is Diversity 392 Integrates Multiple Hunger
Signals 433
12.4 F
 or Many Vertebrates, Parental Care
Determines Offspring Survival 396 The Cutting Edge ■ Friends with
Benefits 440
SEXUAL DIFFERENTIATION 397
13.6 O
 besity and Eating Disorders Are
12.5 S
 ex Determination and Sexual Difficult to Treat 441
Differentiation Occur Early in
BOX 13.2 Body Fat Stores Are Tightly
Development 397 Regulated, Even after Surgical Removal
12.6 G
 onadal Hormones Direct Sexual of Fat 442
Differentiation of the Brain and Visual Summary 447
Behavior 402

14
BOX 12.1 The Paradoxical Sexual
Differentiation of the Spotted Hyena 406
 iological Rhythms, Sleep,
B
and Dreaming 449
12.7 D
 o Fetal Hormones Masculinize
Human Behaviors in When Sleep Gets Out of Control 449
Adulthood? 410
BIOLOGICAL RHYTHMS 450
Vi su a l S umm a r y 41 5
14.1 Many Animals Show Daily Rhythms in
Activity 450

13 Homeostasis 417
Active Regulation of the Internal
Environment
14.2 T
 he Hypothalamus Houses a
Circadian Clock 451
SLEEPING AND WAKING 457
Harsh Reality TV 417
14.3 H
 uman Sleep Exhibits Different
13.1 H
 omeostasis Maintains a Consistent Stages 457
Internal Environment: The Example
of Thermoregulation 418 14.4 W
 hy and How Did Sleep
BOX 13.1 Physiological and Behavioral Evolve? 463
Thermoregulation Are Integrated 422 BOX 14.1 Sleep Deprivation Can Be Fatal 464
FLUID REGULATION 422 14.5 A
 t Least Four Interacting Neural
Systems Underlie Sleep 471
13.2 W
 ater Shuttles between Two Body
Compartments 422 The Cutting Edge ■ Can Individual Neurons
Be “Sleepy”? 476
13.3 T
 wo Internal Cues Trigger
Thirst 425 14.6 S
 leep Disorders Can Be Serious,
Even Life-Threatening 478
FOOD AND ENERGY Visual Summary 482
REGULATION 429

CONTENTS XI
PART V Emotions and Mental Disorders 485

15  motions, Aggression,
E
and Stress 487 16 Psychopathology 521
Biological Basis of Behavioral
Disorders
The Hazards of Fearlessness 487
“The Voice” 521
15.1 B
 road Theories of Emotion
Emphasize Bodily Responses 488 16.1 S
 chizophrenia Is the Major
Neurobiological Challenge in
BOX 15.1 Lie Detector? 491 Psychiatry 523
15.2 D
 id a Core Set of Emotions Evolve BOX 16.1 Long-Term Effects of
in Humans and Other Animals? 492 Antipsychotic Drugs 532

15.3 S
 pecialized Neural Mechanisms 16.2 D
 epression Is the Most Prevalent
Mediate the Experience and Mood Disorder 537
Expression of Emotions 496 The Cutting Edge ■ Can Our Genes Tell Us
The Cutting Edge ■ Synaptic Changes Which Drugs to Use? 541
during Fear Conditioning 502
BOX 16.2 The Season to Be Depressed? 545
15.4 N
 eural Circuitry, Hormones, and
Synaptic Transmitters Mediate 16.3 E
 xtreme Mood Cycles Define
Violence and Aggression 506 Bipolar Disorder 545

15.5 S
 tress Activates Many Bodily 16.4 T
 here Are Several Types of Anxiety
Responses 510 Disorders 547
Visu a l S um m a r y 51 9 BOX 16.3 Tics, Twitches, and Snorts:
The Unusual Character of Tourette's
Syndrome 552
Visual Summary 553

PART VI Cognitive Neuroscience 555

17 
Learning and Memory 557 BOX 17.1 Emotions and Memory

NEURAL MECHANISMS OF MEMORY


575

Trapped in the Eternal Now 557 STORAGE 576


FUNCTIONAL PERSPECTIVES ON 17.4 M
 emory Storage Requires Physical
LEARNING AND MEMORY 558 Changes in the Brain 576

17.1 There Are Several Kinds of Learning 17.5 S


 ynaptic Plasticity Can Be Measured
and Memory 558 in Simple Hippocampal Circuits 582
The Cutting Edge ■ Artificial Activation
17.2 D
 ifferent Forms of Nondeclarative
of an Engram 587
Memory Involve Different Brain
Regions 565 17.6 In the Adult Brain, Newly Born
Neurons May Aid Learning 589
17.3 S
 uccessive Processes Capture, Store,
and Retrieve Information in the Visual Summary 593
Brain 569

XII CO NT E NTS
18  ttention and Higher
A 19.3 L
 eft-Hemisphere Damage Can
Cognition 595 Cause Aphasia 641

One Thing at a Time 595 19.4 C


 ompeting Models Describe
the Left-Hemisphere Language
ATTENTION 596 System 645
18.1 A
 ttention Selects Stimuli for VERBAL BEHAVIOR: SPEECH AND
Processing 596 READING 651
BOX 18.1 Reaction Time Responses, 19.5 L
 anguage Has Both Learned and
from Input to Output 600 Unlearned Components 652
18.2 Targets of Attention: Attention BOX 19.2 Williams Syndrome Offers Clues
Alters the Functioning of Many about Language 654
Brain Regions 603 BOX 19.3 Vocal Behavior in Birds and
18.3 Sources of Attention: A Network Other Species 657
of Brain Sites Creates and Directs 19.6 R
 eading Skills Are Difficult
Attention 609 to Acquire and Frequently
CONSCIOUSNESS AND EXECUTIVE Impaired 658
FUNCTION 616 RECOVERY OF FUNCTION AFTER
18.4 Consciousness Is a Mysterious BRAIN DAMAGE 662
Product of the Brain 616 19.7 S
 tabilization and Reorganization
BOX 18.2 Phineas Gage 623 Are Crucial for Recovery of
Function 662
The Cutting Edge ■ Building a Better
Mind Reader 627 BOX 19.4 The Amazing Resilience of a
Child’s Brain 663
Vi su a l S umm a r y 6 29
The Cutting Edge ■ Contact Sports Can
Be Costly 665

19  anguage and
L
Lateralization 631
Visual Summary 667

Silencing the Inner Voice 631


Appendix A–1
BRAIN ASYMMETRY AND
LATERALIZATION OF Glossary G–1
FUNCTION 632
References R–1
19.1 T
 he Left and Right Hemispheres
Are Different 632 Author Index AI–1
BOX 19.1 The Wada Test 638
Subject Index SI–1
19.2 R
 ight-Hemisphere Damage Impairs
Specific Types of Cognition 639

CONTENTS XIII
Preface
Twenty-four years ago, a new kind of textbook was published for University courses
that were often called “Brain and Behavior.” As the field evolved, the book’s title
metamorphosed from Biological Psychology to Behavioral Neuroscience, but the same
drive to provide a definitive and comprehensive survey of the neurosciences lies at
the heart of all our efforts. We strive to keep the book up-to-date while keeping a
Courtesy of Dr. Sarah Moghadam, VA Palo Alto Health conversational tone to make this wealth of information not just accessible, but fas-
Care System, Palo Alto, CA and Dr. Ahmad Salehi, Dept. of
Psychiatry & Behavioral Sciences, Stanford Medical School cinating. The biggest change in this new edition is the development of Learning
Objectives for each segment of the book, with the idea that telegraphing what’s to
come will focus readers’ attention and facilitate learning. As you finish each section
of text, it would be a good idea to go back and read the associated Learning Objec-
tives to see whether in fact you incorporated the material. If not, a quick review of
that text may be in order.
As always, there have been plenty of new findings to add to this edition. In fact,
the problem we face is which of the many, many new findings to leave out—those
that are not quite essential for a survey of the field. We are pretty picky about what
we add, and still it seems like a deluge of new information and ideas. Hundreds
of new papers are cited in this edition. If that sounds like a lot, let us give you a
perspective on how many new papers were omitted. On our newsfeed site (www.
biopsychology.com/news/), 1,299 new links were added in 2018 alone. Those are just
the findings that were important enough to get the attention of mass media report-
ers. As we note in Chapter 1, over 40,000 new articles indexed under “neuroscience”
appeared that year in PubMed. It would take a thick tome just to list the titles of the
papers from 2018!
While being very, very selective in sampling this flood of findings, we have made
substantial changes in every chapter. For example, in Chapter 3 we have a new
figure comparing “kiss and run” synapses with more traditional models of synaptic
transmission. Chapter 5 has new material about a hormone secreted from bone that
acts on the hypothalamus to reduce appetite. We totally reorganized Chapter 7 for a
more streamlined approach and discuss the growing doubts about whether amyloid
deposits cause Alzheimer’s. Chapter 9 needed a new figure comparing transduction
in the five taste receptors. Chapter 13 talks about yet another factor affecting appetite,
glucagon-like peptide 1. Chapter 16 now discusses the logic of pharmacogenomics
to treat depression, while Chapter 18 was thoroughly reorganized and includes more
about executive function. Honestly, we could go on like this for every chapter. Clearly
this is an exciting era in the neurosciences. As Lewis Carrol put it, “We must run as
fast as we can just to stay in place!”
We’ve also kept several very popular features from previous editions: The Cutting
Edge appears in each chapter, where we explore some of the most exciting examples
of recent research, and each chapter ends with a Visual Summary, where you can see
graphic reminders as you review the principle findings that we just presented. These
Visual Summaries really shine online, where with just a click you can review figures,
animations, and quizzes to help integrate the material. We also continue to open
each chapter with a gripping vignette, relating someone’s real-life experiences that
will be better understood as the content of the chapter unfolds, and we again replaced
several of these vignettes as more recent events bring to the surface many of the
important issues in behavioral neuroscience. Likewise we’ve retained the marginal
glossary that makes it easy to find the definitions that unlock the material, as well
as two features to let you burrow in on a particular subject: the online supplements

XIV C HAP T E R 
called A Step Further cited throughout the text, and the Recommended Reading
at the close of each chapter.
You might think that approaching the quarter-century mark we’d be jaded about
improving and revising our presentations, but we still love it, perhaps because the
dynamic and exciting pace of neuroscience research shows no sign of abating soon.
As always, we welcome all feedback, praise or criticism, cuts or additions, from our
readers. You can email us directly at behavneuro@gmail.com.

Acknowledgments
We continue to feel so lucky to work with the inestimable team at Sinauer Associates,
now a part of Oxford University Press, whose deep skills and generous guidance
transform our hundreds of files, thousands of email attachments and sometimes
scrambled emails into yet another beautiful book. Again, we feel so grateful to benefit
from the experience and exquisite taste of others. In particular, the book could not
exist without the contributions of Senior Acquisition Editor Syd Carroll, Production
Editor Alison Hornbeck, Production Manager Joan Gemme, Book Designer and
Production Specialist Annette Rapier, and Media and Supplements Editor Zan Carter
and her crew. We also fondly bid adieu to the recently retired Chris Small, Production
Manager for all our previous editions. We hope you’re enjoying yourself, Chris, but
how could you abandon us!? A cadre of commandos delved deep in the archives to
deal with copyrights and permissions, so we salute you Michele Beckta, Mark Siddall,
and Tracy Marton. We’d also like to thank our copy editor Lou Doucette, and our
longtime art studio, Dragonfly Media, who bring amazing skill and commitment to
make us look good.
We must also thank the founder of Sinauer Associates, Andy Sinauer, for his
unwavering support over the years, with a touch of sadness upon his retirement.
We are so proud to be a part of Andy’s tremendous legacy, begun all those years
ago with From Neuron to Brain, creating gorgeous books that make even the most
complex topics accessible and enjoyable.
By this point in the evolution of the book, we have benefited from the wisdom
and advice of hundreds of colleagues who have generously served as reviewers of
past editions. Although we don’t have the space to list them all, we want to ac-
knowledge that in many ways the book you are holding is the product of a whole
community of neuroscientists. In this, the Ninth Edition, the following colleagues
have provided invaluable critique and commentary:

Susan Bachus, University of Maryland, Baltimore County


Susan Barron, University of Kentucky
Christopher Beeman, Central Washington University
Jin Bo, Eastern Michigan University
David Brodbeck, Algoma University
Elizabeth Caldwell, University of New Hampshire
James Cherry, Boston University
Michael Cohen, Loyola University Chicago
Paul J. Currie, Reed College
Patrick Cushen, Murray State University
Deana Davalos, Colorado State University
Darragh P. Devine, University of Florida
Christopher W. Drapeau, Valparaiso University
Kelli A. Duncan, Vassar College
Raymond H. Dye, Jr., Loyola University Chicago
Taffeta Elliott, New Mexico Institute of Mining and Technology
Alison A. Fedio, Argosy University, Northern Virginia
Sara B. Festini, University of Tampa

P R EFACE XV
Cynthia Michelle Finley, College of Marin
Jonathan Franz, SUNY Empire State College
Koren Ganas, University of Illinois
Sophie George, Dixie State University
Aaron Godlaski, Centre College
Brian J. Hock, Austin Peay State University
Jennifer Ingemi, Northeastern University
Mary Ellen Kelly, Haverford College
Susan Kennedy, Denison University
Michael Kerchner, Washington College
Sarita Lagalwar, Skidmore College
Stephen Lippi, George Mason University
Mario L. Mata, California State University, Los Angeles
Alexandra Roach, University of South Carolina, Aiken
Russell Romeo, Barnard College of Columbia University
Timothy Roth, Franklin and Marshall College
Emma Sarro, Dominican College
Peter A. Serrano, Hunter College, City University of New York
Fredric Shaffer, Truman State University
KatieAnn Skogsberg, Centre College
Lucy J. Troup, University of the West of Scotland
Adriana Uruena-Agnes, St. Petersburg College
Jennifer Wilhelm, College of Charleston
Jan R. Wessel, University of Iowa
Susan Zup, University of Massachusetts Boston

Finally, we thank all those tireless colleagues trying to understand the neural basis of
behavior, with techniques that would have seemed like sorcery only a few years ago,
and who share their hard-won findings with us all.

S. Marc Breedlove Neil V. Watson

XVI PREFACE
Media and
Supplements
to accompany
Behavioral Neuroscience,
Ninth Edition
Courtesy of Dr. Sarah Moghadam, VA Palo Alto Health
Care System, Palo Alto, CA and Dr. Ahmad Salehi, Dept. of
Psychiatry & Behavioral Sciences, Stanford Medical School

For the Student


Companion Website (bn9e.com)
The Behavioral Neuroscience Companion Website contains a range of study and review
resources to help students master the material presented in each chapter of the text-
book. Access to the site is included with each new copy of the textbook (see inside
front cover). The site includes the following resources:
• Chapter Outlines that outline each chapter and link to relevant Study Questions
• Brain Explorer that offers an interactive way to explore the brain anatomy discussed
in each chapter
• Activities that help the student review key structures and processes
• Animations and Videos that illustrate many of the complex, dynamic concepts
and processes of behavioral neuroscience
• Media Clips that highlight interesting topics in the chapters (NEW for this edition)
• “A Step Further” essays that offer expanded coverage of selected topics
• Visual Summaries that link to all the Activities, Animations, and Videos,
forming a complete review of each chapter
• Study Questions that help the student master the full range of material in each chapter
• Flashcards that review and reinforce the many new terms introduced in each chapter
• Complete Glossary that provides quick access to definitions of all the important
terminology in the textbook

BioPsychology NewsLink (bn9e.com/news)


This invaluable online resource helps students make connections between the science
of behavioral neuroscience and their daily lives and keeps them apprised of the lat-
est developments in the field. The site includes links to thousands of news stories, all
organized both by keyword and by textbook chapter. The site is updated 3–4 times per
week, so it includes up-to-the-minute information. NewsLink updates are also avail-
able on Facebook (facebook.com/behavioralneuroscience).
For the Instructor
Ancillary Resource Center (oup-arc.com)
The Ancillary Resource Center (ARC) provides instructors using Behavioral Neurosci-
ence 9e with a wide variety of resources to aid in course planning, lecture development,
and student assessment. Content includes:
• Figures & Tables: All the figures, photos, and tables from the textbook are provided as
JPEGs, all optimized for use in presentations.
• PowerPoint Presentations: Two PowerPoint presentations are provided for each
chapter of the textbook:
• Figures: All the chapter’s figures, photos, and tables, with titles and complete
captions
• Lectures: Complete lecture outlines, including selected figures
• Instructor’s Manual: The Instructor’s Manual includes useful resources for planning
your course, lectures, and exams. For each chapter of the textbook, the IM includes
a chapter overview, a chapter outline, the chapter’s key concepts, additional refer-
ences for course and lecture development, and a list of the chapter’s key terms.
• Videos: A robust collection of video segments from the BBC and other sources
bring to life many important concepts discussed in the textbook. These videos
can be used as excellent lecture-starters and/or discussion topics.
• Animations: These detailed animations from the Companion Website help
enliven lectures and illustrate dynamic processes.
• Animation Quizzes: These quizzes test the student’s understanding of the topic
(NEW for this edition).
• Chapter Quizzes: Quiz questions for each chapter in two formats: Available
in Blackboard, Canvas, D2L platform, or as MS Word files.
• Multiple choice tests student comprehension of the material covered in
each chapter.
• Essays challenge students to synthesize and apply what they have learned.
• Test Bank: The Test Bank consists of a broad range of questions covering key facts
and concepts in each chapter. Multiple choice, essay, and paragraph development
questions are included. Questions are ranked according to Bloom’s Taxonomy and
referenced to specific textbook sections. NEW for this edition, questions are also
aligned to the textbook Learning Objectives. (Available in Blackboard, Canvas,
D2L platform, or as MS Word files.)

Interoperable Course Cartridge


At Oxford University Press, we create high quality, engaging, and affordable digital
material in a variety of formats, and deliver it to you in the way that best suits the
needs of you, your students, and your institution. With Interoperable Course Cartridge
by Oxford University Press, there is no need for you and your students to learn a sepa-
rate publisher-provided courseware platform in order access quality digital learning
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content from OUP directly into their LMS for assigning and grading.

XVIII M E DI A AN D S U P P L E ME N TS 
Value Options
eBook
(ISBN 978-1-60535-937-3)
Ideal for self-study, the Behavioral Neuroscience, Ninth Edition, enhanced eBook deliv-
ers the full suite of digital resources in a format that is independent from any course-
ware or learning management system platform. The enhanced eBook is available
through leading higher education eBook vendors.

Looseleaf Textbook
(ISBN 978-1-60535-936-6)
Behavioral Neuroscience, Ninth Edition is also available in a three-hole-punched,
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integrate instructor material with the text.

ME DIA AND SUP P LEMENTS  XIX


Psychiatry & Behavioral Sciences, Stanford Medical School
Care System, Palo Alto, CA and Dr. Ahmad Salehi, Dept. of
Introduction

Courtesy of Dr. Sarah Moghadam, VA Palo Alto Health


Scope and Outlook
Machine or Human?
In the near future depicted in the HBO series Westworld, people visit a theme park set
in the Old West, with steam locomotives, saloons, and brothels, populated with an-
droids, called “hosts,” to entertain humans. The mechanical hosts provide their guests
with anything, from casual banter to gunfights, harmless flirting to kinky sex, the only

1
restriction being that the robots are never to harm the humans. The android hosts are
so lifelike in appearance and behavior that visitors may have a hard time distinguishing
whether someone is a fellow guest or a robot. To make the androids’ simulation of hu-
mans complete, they are given backstories, false memories of a life before their appear-
ance for each new batch of guests. Importantly, none of the androids know that they
are mechanical beings rather than humans. It’s probably not much of a spoiler to say
that several plot lines in the series hinge on androids slowly discovering their true nature,
moving from shock and shame that they are mere machines, to openly rebelling from
the notion that they are to be used, and abused, as mere playthings for the humans.
We aren’t told too much about how the android “brains” in Westworld work, be-
cause, of course, such technology remains far outside our grasp, so the writers,
reduced to mere speculation, remain rather vague. But apparently the knowledge and
personality for any particular android lies in a “control unit,” a golf-ball-size device that
can be extracted from the head of one host and implanted into the head of another,
interchangeable body. Presumably, if we had enough knowledge and surgical skill, we
could remove your brain from your head and connect it up to the head of some other
body. Would you still be you? Even if we put your brain into a body of the opposite sex?
Come to think of it, are you entirely sure there is a brain in your head, and not one of
those control units?

Our aim in this book is to help you learn what is known so far about how brains work,
and about how much more we have yet to learn. We will explore the many ways in
which the structures and actions of the brain produce mind and behavior. But that is
only half of our task. We are also interested in the ways in which behavior and experi-
ence modify the structures and actions of the brain. One of the most important lessons
we want to convey is that interactions between brain and behavior are reciprocal. The
brain controls behavior and, in turn, behavior and experience alter the brain.
We hope to give an interesting account of the main ideas and research in be-
havioral neuroscience, which is of great popular as well as scientific interest. Most
important, we try to communicate our own interest and excitement about the mys-
teries of mind and body.

Go to Brain Explorer
bn9e.com/be1
1.1 The Brain Is Full of Surprises
Learning Objectives
After reading this section, you should be able to:
1.1.1 
Name the main type of cells found in the brain, and name the connections
between them.
1.1.2 
List the names of some of the many fields of study related to behavioral
neuroscience.
1.1.3 
Describe five different perspectives taken in understanding the biology
of behavior.

I used to think that the brain was the most wonderful organ in my body. Then I
realized who was telling me this.
—Emo Philips
(American comedian)

Of course we should always consider the source when evaluating an idea, but even
so, the brain indeed seems like a pretty wonderful organ. For one thing, brains pro-
duced the entire extent of human knowledge, everything we understand about the
universe, however limited that may be. Brains also produced every written descrip-
tion of that hard-won knowledge (including this book you hold in your hands), as
well as every work of visual art, from doodles to the sweeping frescos on the ceiling
of the Sistine Chapel.
Most of us have a hard time grasping the idea of a billion of anything, but your
head contains an estimated 86 billion nerve cells, or neurons (from the Greek word
for “nerve” or “cord”) (Herculano-Houzel, 2012). Each neuron contacts many other
cells at points called synapses, so there are trillions of those between your ears. A
specialized extension of neurons, called an axon, is microscopically slender, yet it may
neuron Also called nerve cell. The basic be several feet long. We’ll learn that axons produce electrical impulses that travel
unit of the nervous system. hundreds of miles per hour. FIGURE 1.1 offers a list of just a few of the things we will

© Dwayne Godwin, 2011

1.1 Your Brain by the Numbers The cerebral cortex is the outermost portion of the brain.

2 C HA PT E R 1
learn about the human brain in the course of this book. All this hardware isn’t just for neuroscience The study of the
show—it allows you to take in all the information in that figure in less than a minute. nervous system.
behavioral neuroscience Also called
What is behavioral neuroscience? biological psychology. The study of the
No treaty or trade union agreement defines the boundaries of behavioral neuroscience. neural bases of behavior and mental
The first people to study the relationships between brain and behavior regarded them- processes.
selves as philosophers, and their findings contributed to the births of biology and psy-
chology. Those disciplines merged in the twentieth century to form biological psychology,
the field that relates behavior to bodily processes. With the modern explosion of neuro-
science, the study of the brain, this research has evolved to the point that behavioral
neuroscience offers a more accurate description. Whichever name is used, the main
goal of this field is to understand the neuroscience underlying behavior and experience.
Behavioral neuroscience is a field that includes many players who come from quite
different backgrounds: psychologists, biologists, physiologists, engineers, neurolo-
gists, psychiatrists, and many others. Thus, there are many career opportunities, in
both universities and private industry, for people with interests in this field (Hitt,
2007). FIGURE 1.2 maps the relations of behavioral neuroscience to these many oth-
er disciplines. Clearly, the behavioral neuroscience umbrella opens very wide.

Cognitive
science
Computer
Anthropology
science

Cognitive
psychology
Evolutionary Sociobiology Artificial Psychiatry
biology intelligence
Cognitive
Behavioral neuroscience Behavioral
ecology/ethology Social Neural medicine
neuroscience modeling
Comparative/ Health
Paleontology evolutionary psychology Neurology
Paleoneuro- psychology Clinical
Cognitive neuro-
anatomy Comparative
neuro- psychology
neuroanatomy BEHAVIORAL psychology
Neural NEUROSCIENCE Neuro-
Neuro- imaging physiology Electro-
anatomy physiology
Anatomy Developmental Psycho- Physiology
psychobiology pharmacology
Behavior Psychoneuro-
Developmental genetics immunology Pharmacology
neurobiology Behavioral
endocrinology
Developmental Genetics/ Neuro- Biochemistry
biology epigenetics immunology
Neuro-
endocrinology

Molecular Immunology
biology
Endocrinology

1.2 What’s in a Name? In this graphical representation of the relationships among behavioral
neuroscience and other scientific disciplines, fields toward the center of the map are closest to
behavioral neuroscience in their history, outlook, aims, and/or methods.

In trodu ctio n 3
Five viewpoints explore the biology of behavior
In our effort to understand the neuroscience bases of behavior, we use several dif-
ferent perspectives. Because each one yields information that complements the
others, the combination of perspectives is especially powerful. We will discuss five
major perspectives:
1. Describing behavior
2. Observing the development of behavior and its biological characteristics
over the life-span
3. Studying the biological mechanisms of behavior
4. Studying applications of behavioral neuroscience—for example, its application
to dysfunctions of human behavior
5. Studying the evolution of behavior

These perspectives are discussed in the sections that follow, and TABLE 1.1 illustrates
how each perspective can be applied to three kinds of behavior.

Behavior can be described according to different criteria


Until we describe what we want to study, we cannot accomplish much. Depending
on our goals, we may describe behavior in terms of detailed acts or processes, or in
terms of results or functions. An analytical description of arm movements might
record the successive positions of the limb or the contraction of different muscles.
A functional behavioral description, by contrast, would state whether the limb was
being used in walking or running, texting or sexting. To be useful for scientific
study, a description must be precise and reveal the essential features of the behavior,
using accurately defined terms and units.

TABLE 1.1 Five Research Perspectives Applied to Three Kinds of Behavior


Language and
Research perspective Sexual behavior Learning and memory communication
DESCRIPTION
Structural What are the main patterns In what main ways does How are the sounds of speech
of reproductive behavior behavior change as a patterned?
and sex differences in consequence of experience—
behavior? for example, conditioning?
Functional How do specialized patterns How do certain behaviors lead What behavior is involved in
of behavior contribute to rewards or avoidance of making statements or asking
to mating and to care of punishment? questions?
young?
ONTOGENY How do reproductive How do learning and memory What changes in the brain
(development) and secondary sex change as we grow older? when a child learns to
characteristics develop speak?
over the life-span?
MECHANISMS What neural circuits and What anatomical and chemical What brain regions are
hormones are involved in changes in the brain hold particularly involved in
reproductive behavior? memories? language?
APPLICATIONS Low doses of testosterone Gene therapy and behavioral Speech therapy, in conjunction
restore libido in some therapy improve memory in with amphetamine treatment,
postmenopausal women. some senile patients. speeds language recovery
following stroke.
EVOLUTION How does mating depend How do different species How did the human speech
on hormones in different compare in kinds and speed apparatus evolve?
species? of learning?

4 C HA PT E R 1
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FINE ALMOND CAKE.

Blanch, dry, and pound to the finest possible paste, eight ounces
of fresh Jordan almonds, and one ounce of bitter; moisten them with
a few drops of cold water or white of egg, to prevent their oiling; then
mix with them very gradually twelve fresh eggs which have been
whisked until they are exceedingly light; throw in by degrees one
pound of fine, dry, sifted sugar, and keep the mixture light by
constant beating, with a large wooden spoon, as the separate
ingredients are added. Mix in by degrees three-quarters of a pound
of dried and sifted flour of the best quality; then pour gently from the
sediment a pound of butter which has been just melted, but not
allowed to become hot, and beat it very gradually, but very
thoroughly, into the cake, letting one portion entirely disappear
before another is thrown in; add the rasped or finely-grated rinds of
two sound fresh lemons, fill a thickly-buttered mould rather more
than half full with the mixture, and bake the cake from an hour and a
half to two hours in a well-heated oven. Lay paper over the top when
it is sufficiently coloured, and guard carefully against its being
burned.
Jordan almonds, 1/2 lb.; bitter almonds, 1 oz.; eggs, 12; sugar, 1
lb.; flour, 3/4 lb.; butter, 1 lb.; rinds lemons, 2: 1-1/2 to 2 hours.
Obs.—Three-quarters of a pound of almonds may be mixed with
this cake when so large a portion of them is liked, but an additional
ounce or two of sugar, and one egg or more, will then be required.
PLAIN POUND OR CURRANT CAKE.

(Or rich Brawn Brack, or Borrow Brack.)


Mix, as directed in the foregoing receipt, ten eggs (some cooks
take a pound in weight of these), one pound of sugar, one of flour,
and as much of butter. For a plum-cake, let the butter be worked to a
cream; add the sugar to it first, then the yolks of the eggs, next stir
lightly in the whites, after which, add one pound of currants and the
candied peel, and, last of all, the flour by degrees, and a glass of
brandy when it is liked. Nearly or quite two hours’ baking will be
required for this, and one hour for half the quantity.
To convert the above into the popular Irish “speckled bread,” or
Brawn Brack of the richer kind, add to it three ounces of carraway-
seeds: these are sometimes used in combination with the currants,
but more commonly without. To ice a cake see the receipt for Sugar
Glazings at the commencement of this Chapter, page 543. A rose-
tint may be given to the icing with a little prepared cochineal, as we
have said there.
RICE CAKE.

Take six eggs, with their weight in fine sugar, and in butter also,
and half their weight of flour of rice, and half of wheaten flour; make
the cake as directed for the Madeira or almond cake, but throw in the
rice after the flour; then add the butter in the usual way, and bake the
cake about an hour and ten minutes. Give any flavour that is liked.
The butter may be altogether omitted. This is a moderate-sized cake.
Eggs, in the shell, 6; their weight in butter and in sugar; half as
much flour of rice, and the same of wheaten flour: 1 hour, 10
minutes.
WHITE CAKE.

Beat half a pound of fresh butter to a cream, add to it an equal


weight of dried and sifted sugar, the yolks and whites of eight eggs,
separately whisked, two ounces of candied orange-peel, half a
teaspoonful of mace, a glass of brandy, one pound of flour strewed
in by degrees, and last of all a pound and a quarter of currants.
Directly it is mixed send the cake to a well-heated oven, and bake it
for two hours. Four ounces of pounded almonds are sometimes
added to it.
Butter, 1/2 lb.; sugar, 1/2 lb.; eggs, 8; mace, 1/2 teaspoonful;
brandy, 1 wineglassful; flour, 1 lb.; candied-peel, 2 oz.; currants, 1-
1/2 lb.: 2 hours.
A GOOD SPONGE CAKE.

Rasp on some lumps of well-refined sugar the rind of a fine sound


lemon, and scrape off the part which has imbibed the essence, or
crush the lumps to powder, and add them to as much more as will
make up the weight of eight or ten fresh eggs in the shell; break
these one by one, and separate the whites from the yolks; beat the
latter in a large bowl for ten minutes, then strew in the sugar
gradually, and beat them well together. In the mean time let the
whites be whisked to a quite solid froth, add them to the yolks, and
when they are well blended sift and stir the flour gently to them, but
do not beat it into the mixture; pour the cake into a well-buttered
mould, and bake it an hour and a quarter in a moderate oven.
Rasped rind, 1 large lemon; fresh eggs, 8 or 10; their weight of
dry, sifted sugar; and half their weight of flour: baked, 1-1/4 hour,
moderate oven.
A SMALLER SPONGE CAKE.

(Very good.)
Five full-sized eggs, the weight of four in sugar, and of nearly three
in flour, will make an exceedingly good cake: it may be flavoured, like
the preceding one, with lemon-rind, or with bitter almonds, vanilla, or
confected orange-blossoms reduced to powder. An hour will bake it
thoroughly. All the ingredients for sponge cakes should be of good
quality, and the sugar and flour should be dry; they should also be
passed through a fine sieve kept expressly for such purposes. The
excellence of the whole depends much on the manner in which the
eggs are whisked: this should be done as lightly as possible, but it is
a mistake to suppose that they cannot be too long beaten, as after
they are brought to a state of perfect firmness they are injured by a
continuation of the whisking, and will at times curdle, and render a
cake heavy from this cause.
FINE VENETIAN CAKE OR CAKES.

Take of sound Jordan almonds, blanched and well dried at the


mouth of a cool oven or in a sunny window, seven ounces, full
weight, and one of bitter almonds with them; pound the whole to a
perfect paste with a few drops of white of egg or orange-flower
water; then mix them thoroughly with one pound of flour and eight
ounces of butter (which should be cool and firm, or it will render the
paste too soft), and break this down quite small; then add eight
ounces of pounded sugar, on part of which the rind of a fine lemon
has been rasped previously to its being crushed to powder. Make
these into a paste with the yolks of four eggs, or with rather less
should they be large, for if too moist, it will adhere to the board and
roller. To make a Venetian cake of moderate size, roll the paste less
than a quarter of an inch thick, and cut with the larger fluted cutter,
shown at page 376, six or seven portions of equal size; lay them on
lightly floured or buttered tins, and bake them in a slow oven until
they are firm and crisp, and equally coloured of a pale brown. Should
they seem to require it, lay them one on the other, while they are still
warm, and place a baking-tin, with a slight weight upon them to
render them quite level. When they are cold, spread upon each a
different kind of choice preserve, and pile the whole evenly into the
form of an entire cake. The top may be iced, and decorated with
pistachio-nuts, or grains of coloured sugar, or with a wreath of
almond-paste leaves. There are many varieties of this dish, which is
known by different names in different countries. It is sometimes
called a Neapolitan Cake, sometimes a Thousand Leaf Cake à la
Française. It is occasionally made entirely of almond-paste, and
highly decorated; it may be formed also of many layers of puff or fine
short crust cut of uniform size, or gradually less, so as to leave round
each a clear border of an inch wide, which may be covered with
coloured icing, or ornamented with preserved fruit, tinted almonds,
grains of white or pink sugar candy, or aught else that the fancy may
direct.
To make the small Venetian cakes, roll the paste directed for the
large one at the commencement of this receipt, into balls, flatten
them with the hand to about the third of an inch thick, brush them
with beaten egg, and cover them plentifully with white sugar-candy
crushed about half the size of a pea: bake them in a slow oven.
Almonds, 8 oz.; flour, 1 lb.; butter, 8 oz.; sugar, 1/2 lb.; rind of 1
lemon; yolks of eggs, 3 to 4; preserve as needed.
A GOOD MADEIRA CAKE.

Whisk four fresh eggs until they are as light as possible, then,
continuing still to whisk them, throw in by slow degrees the following
ingredients in the order in which they are written: six ounces of dry,
pounded, and sifted sugar; six of flour, also dried and sifted; four
ounces of butter just dissolved, but not heated; the rind of a fresh
lemon; and the instant before the cake is moulded, beat well in the
third of a teaspoonful of carbonate of soda: bake it an hour in a
moderate oven. In this, as in all compositions of the same nature,
observe particularly that each portion of butter must be beaten into
the mixture until no appearance of it remains before the next is
added; and if this be done, and the preparation be kept light by
constant and light whisking, the cake will be as good, if not better,
than if the butter were creamed. Candied citron can be added to the
paste, but it is not needed.
Eggs, 4; sugar, 6 oz.; flour, 6 oz.; butter, 4 oz.; rind of 1 lemon;
carbonate of soda, 1/3 of teaspoonful: 1 hour, moderate oven.
A SOLIMEMNE.

A rich French breakfast cake, or Sally Lunn.


From three-quarters of a pound of flour take three ounces for the
leaven, and make it into a lithe paste with half an ounce of solid,
well-washed yeast (see Chapter XXXI.), mixed with two or three
tablespoonsful of just warm cream, or new milk; throw a cloth over
and leave it near the fire to rise for about half an hour, or until it is
twice its original size. In the interim make a hollow in the centre of
the remainder of the flour, and put into it a quarter of an ounce of fine
salt, one ounce of pounded sugar, the yolks of four fresh eggs, four
ounces of lukewarm butter, and a couple of tablespoonsful of cream,
also warm. Mix the whole gently and carefully into a perfectly smooth
paste, flatten it with the hand upon the dresser, spread the leaven
over it, and blend them thoroughly with light kneading, as directed for
brioche paste, page 349. The whole should be of the same colour
throughout.
Next, put it into a small, well-buttered copper stewpan, or plain
cake-mould, and let it remain in a moderately warm place until it has
risen, like the leaven, to double its original size; then with a paste-
brush or feather wash the top with beaten egg, and without
disturbing it, set it into a tolerably quick oven, and bake it nearly or
quite an hour; but do not allow it to be too deeply coloured. Turn it
from the mould, cut it once or twice asunder, and pour over the slices
plenty of good butter, just dissolved in a small saucepan; put the
cake together again, and serve it immediately. It may be converted
into an excellent entremets by spreading currant, or other fine jelly,
or preserve, quickly upon it when it is cut, and sifting sugar thickly on
the top after it is restored to its proper form: it is then called a
Dresden cake. We think that when left until cold and toasted, the
solimemne is even better than when served hot. It will be many
hours rising; sometimes as many as six or eight. If wanted for
breakfast it should be made over night.
Flour 3/4 lb.; yeast, 1/2 oz.; little cream; salt, 1/4 oz.; sugar, 1 oz.;
yolks of eggs, 4; butter, 4 oz.: to rise from 6 to 8 hours. Baked 1
hour.
BANBURY CAKES.

First, mix well together a pound of currants, cleaned with great


nicety and dried, a quarter-pound of beef suet, finely minced, three
ounces each of candied orange and lemon-rind, shred small, a few
grains of salt, a full quarter-ounce of pounded cinnamon and nutmeg
mixed, and four ounces of macaroons or ratafias rolled to powder.
Next, make a light paste with fourteen ounces of butter to the pound
of flour; give it an extra turn or two to prevent its rising too much in
the oven; roll out one half in a very thin square, and spread the
mixed fruit and spice equally upon it; moisten the edges, lay on the
remaining half of the paste, rolled equally thin, press the edges
securely together, mark the whole with the back of a knife in regular
divisions of two inches wide and three in length, bake the pastry in a
well-heated oven from twenty-five to thirty minutes, and divide it into
cakes while it is still warm. They may be served as a second course
dish either hot or cold, and may be glazed at pleasure.
Currants, 1 lb.; beef-suet, 4 oz.; candied orange and lemon-rind
each, 3 oz.; salt, small pinch; mixed spices, 1/4 oz.; macaroons or
ratafias, 4 oz.: baked 25 to 30 minutes.
MERINGUES.

Whisk, to the firmest


possible froth, the whites
of six very fresh eggs,
taking every precaution
against a particle of the
yolk falling in amongst
them. Lay some squares
or long strips of writing-
paper closely upon a
board or upon very clean
trenchers, which ought to
be nearly or quite an inch
thick, to prevent the
meringues from receiving
any colour from the bottom
of the oven. When all is ready, mix with the eggs three-quarters of a
pound of the finest sugar, well dried, and sifted; stir them together for
half a minute, then with a table or dessertspoon lay the mixture
quickly on the papers in the form of a half-egg, sift sugar over them
without delay, blow off with the bellows all that does not adhere, and
set the meringues into a gentle oven. The process must be
expeditious, or the sugar melting will cause the cakes to spread,
instead of retaining the shape of the spoon, as they ought. The
whole art of making them, indeed, appears to us to consist in
preserving their proper form, and the larger the proportion of sugar
worked into the eggs, the more easily this will be done. When they
are coloured to a light brown, and are firm to the touch, draw them
out, turn the papers gently over, separating the meringues from
them, and with a teaspoon scoop out sufficient of the insides to form
a space for some whipped cream or preserve, and put them again
into the oven upon clean sheets of paper, with the moist sides
uppermost, to dry: when they are crisp through they are done. Let
them become cold; fill, and then join them together with a little white
of egg so as to give them the appearance shown in the plate. Spikes
of pistachio nuts, or almonds, can be stuck over them, as
represented there, at pleasure. They afford always, if well made, a
second course dish of elegant appearance, and they are equally
ornamental to breakfasts or suppers of ceremony. They are made in
perfection by the pastry-cooks in France, being equally light,
delicate, and delicious. Much of their excellence, it must be
observed, depends at all times on the attention they receive in the
baking, as well as in the previous preparation. They must, of course,
be quite cold before the preserve or cream is laid into them. From
four to six ounces of almonds, finely powdered, may be smoothly
mixed with the other ingredients for them; and they may be flavoured
with citron, lemon, or orange-rind by rasping the skins of the fruit with
part of the sugar with which they are to be made; then drying, and
reducing it to powder.
Whites of very fresh eggs, 6; sugar, 3/4 lb.: gentle oven, 20 to 30
minutes.
ITALIAN MERINGUES.

Take for these the proportion of whites of eggs and sugar already
indicated in the receipt for Nesselrôde pudding, page 491, that is to
say, six to the pound, or half that quantity for a small number of
meringues. Boil the sugar with a pint of water until it whitens, and
begins to fall in flakes from the skimmer; have the eggs whisked to a
perfectly solid froth quite ready at the proper moment, and when the
sugar has stood for two or three minutes, and been worked well from
the sides of the pan, mingle them gradually, but very quickly, with it,
that the mass may be quite smooth; continue to stir them until they
become firm enough to retain their shape perfectly when moulded
with a teaspoon; lay out the cakes on paper, and place them in an
oven so slow as to harden without giving them colour. As they are
not to be filled, but merely fastened together, they may be baked on
tins. Part of them may be varied by the addition of three or four
ounces of pounded almonds mixed thoroughly with the remainder of
the eggs and sugar, when a portion of the meringues have been
moulded: these, however, will require to be much longer baked than
the others; but they will be excellent. They should be lightly browned,
and crisp quite through.
Sugar, 1 lb.; water, 1 pint; whites of eggs, 6: very slow oven, 20 to
30 minutes, or longer.
THICK, LIGHT GINGERBREAD.

Crumble down very small, eight ounces of butter into a couple of


pounds of flour, then add to, and mix thoroughly with them, half a
pound of good brown sugar, two ounces of powdered ginger, and
half an ounce of ground carraway-seeds; beat gradually to these,
first two pounds of treacle, next three well-whisked eggs, and last of
all half an ounce of carbonate of soda,[174] dissolved in a very small
cupful of warm water; stir the whole briskly together, pour the mixture
into very shallow tins, put it immediately into a moderate oven, and
bake it for an hour and a half. The gingerbread made thus will be
remarkably light and good. For children part of the spice and butter
may be omitted.
174. This should always be of the very best quality when used for cakes.
Carbonate of ammonia is recommended in preference to it by some writers.

Flour, 2 lbs.; butter, 8 oz.; sugar, 1/2 lb.; powdered ginger, 2 oz.;
eggs, 3; carbonate of soda, 1/2 oz.; water, very small cupful: baked
1-1/2 hour.
Obs.—We think that something less than the half ounce of soda
would be sufficient for this gingerbread, for with the whole quantity it
rises in the oven to three times its height, and is apt to run over the
tops of the tins, even when they are but half filled with it at first; or if it
were well beaten into the mass without any water, after being
carefully freed from lumps and mixed with a little sugar, the cake
would still be quite light.
ACTON GINGERBREAD.

Whisk four strained or well-cleared eggs to the lightest possible


froth (French eggs, if really sweet, will answer for the purpose), and
pour to them, by degrees, a pound and a quarter of treacle, still
beating them lightly. Add, in the same manner, six ounces of pale
brown sugar free from lumps, one pound of sifted flour, and six
ounces of good butter, just sufficiently warmed to be liquid, and no
more, for if hot, it would render the cake heavy; it should be poured
in small portions to the mixture, which should be well beaten up with
the back of a wooden spoon as each portion is thrown in: the
success of the cake depends almost entirely on this part of the
process. When properly mingled with the mass, the butter will not be
perceptible on the surface; and if the cake be kept light by constant
whisking, large bubbles will appear in it to the last. When it is so far
ready, add to it one ounce of Jamaica ginger and a large teaspoonful
of cloves in fine powder, with the lightly grated rinds of two fresh full-
sized lemons. Butter thickly, in every part, a shallow square tin pan,
and bake the gingerbread slowly for nearly or quite an hour in a
gentle oven. Let it cool a little before it is turned out, and set it on its
edge until cold, supporting it, if needful, against a large jar or bowl.
We have usually had it baked in an American oven, in a tin less than
two inches deep; and it has been excellent. We retain the name
given to it originally in our own circle.
CHEAP AND VERY GOOD GINGER OVEN-CAKE OR CAKES.

Four French eggs (which must be perfectly sweet, or small English


ones), six ounces of brown sugar of good quality rolled smooth and
fine, six ounces of flour, three of butter, a grain or two of salt, some
grated lemon-rind or candied peel sliced very thin, and half an ounce
or more of ginger in fine powder. Prepare and mix these ingredients
in the order in which they are written, by the directions for “Acton
Gingerbread.” Bake the cake nearly the same time. An American
oven will answer for it perfectly, and it will resemble a really rich
cake, though so cheap. A small quantity of carbonate of soda may
be added quite at last by inexpert cake-makers, to insure its being
light. The same mixture may be baked in small cups or tins in an iron
oven. For a cake of tolerable size half as much again of the
ingredients must be taken, and the whole poured into a round or
square cake-mould.
GOOD COMMON GINGERBREAD.

Work very smoothly six ounces of fresh butter (or some that has
been well washed from the salt, and wrung dry in a cloth) into one
pound of flour, and mix with them thoroughly an ounce of ginger in
fine powder, four ounces of brown sugar, and half a teaspoonful of
beaten cloves and mace. Wet these with three-quarters of a pound
of cold treacle, or rather more, if needful; roll out the paste, cut the
cakes with a round tin cutter, lay them on a floured or buttered
baking tin, and put them into a very slow oven. Lemon-grate or
candied peel can be added, when it is liked.
Flour, 1 lb.; butter, 6 oz.; sugar, 1/4 lb.; ginger, 1 oz.; cloves and
mace, 1/2 teaspoonful; treacle, 3/4 lb.: 1/2 to 3/4 hour.
RICHER GINGERBREAD.

Melt together three-quarters of a pound of treacle and half a pound


of fresh butter, and pour them hot on a pound of flour mixed with half
a pound of sugar and three-quarters of an ounce of ginger. When the
paste is quite cold, roll it out with as much more flour as will prevent
its adhering to the board: bake the cakes in a very gentle oven.

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