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Contents
Preface xvii
1 Principles of Pharmacology 3
Pharmacology: The Science of Drug Action 4
Placebo effect 5
Box 1.1 Pharmacology in Action Naming Drugs 6
Pharmacokinetic Factors Determining Drug
Action 7
Methods of drug administration influence the onset of drug
action 8
Multiple factors modify drug absorption 12
Drug distribution is limited by selective barriers 15
Depot binding alters the magnitude and duration of
drug action 18
Biotransformation and elimination of drugs contribute
to bioavailability 19
Therapeutic Drug Monitoring 25
Box 1.2 Pharmacology in Action Interspecies Drug Dose
Extrapolation 25
2 Structure and Function of the Nervous System 45
Cells of the Nervous System 46
Neurons have three major external features 46
Box 2.1 The Cutting Edge Embryonic Stem Cells 47
Characteristics of the cell membrane are critical for
neuron function 54
Glial cells provide vital support for neurons 55
Box 2.2 Of Special Interest Astrocytes 56
Electrical Transmission within a Neuron 58
Ion distribution is responsible for the cell’s resting
potential 58
Pharmacodynamics: Drug–Receptor
Interactions 27
Box 1.3 Pharmacology in Action Drug Categories 28
Extracellular and intracellular receptors have several
common features 30
Dose–response curves describe receptor activity 31
The therapeutic index calculates drug safety 32
Receptor antagonists compete with agonists for
binding sites 33
Biobehavioral Effects of Chronic Drug Use 34
Repeated drug exposure can cause tolerance 35
Chronic drug use can cause sensitization 38
Pharmacogenetics and Personalized Medicine
in Psychiatry 39
Local potentials are small, transient changes in
membrane potential 60
Sufficient depolarization at the axon hillock opens voltage-
gated Na+ channels, producing an action potential 61
Drugs and poisons alter axon conduction 63
Organization of the Nervous System 65
Box 2.3 The Cutting Edge Finding Your Way in the
Nervous System 65
The nervous system comprises the central and peripheral
divisions 66
CNS functioning is dependent on structural features 68
viii Contents
The CNS has six distinct regions reflecting embryological
development 70
Box 2.4 Of Special Interest Neuroendocrine Response to
Stress 76
3 Chemical Signaling by Neurotransmitters and Hormones 83
Chemical Signaling between Nerve Cells 84
Neurotransmitter Synthesis, Release,
and Inactivation 86
Neurotransmitters encompass several different
kinds of chemical substances 86
Box 3.1 Clinical Applications Orexin-Based
Medications: New Approaches to the Treatment
of Sleep Disorders 87
Neuropeptides are synthesized by a different mechanism
than other transmitters 90
Neuromodulators are chemicals that don’t act like typical
neurotransmitters 91
Classical transmitter release involves exocytosis and
recycling of synaptic vesicles 91
Lipid and gaseous transmitters are not released from
synaptic vesicles 95
Several mechanisms control the rate of neurotransmitter
release by nerve cells 96
Neurotransmitters are inactivated by reuptake and by
enzymatic breakdown 97
4 Methods of Research in Psychopharmacology 117
Research Methods for Evaluating the Brain
and Behavior 118
Techniques in Behavioral Pharmacology 118
Evaluating Animal Behavior 118
Animal testing needs to be valid and reliable to
produce useful information 118
A wide variety of behaviors are evaluated by
psychopharmacologists 120
Box 4.1 Pharmacology in Action Using the Three-
Chamber Social Interaction Test 125
Box 4.2 Clinical Applications Drug Testing for
FDA Approval 131
Techniques in Neuropharmacology 134
The cerebral cortex is divided into four lobes, each
having primary, secondary, and tertiary areas 76
Rat and human brains have many similarities and
some differences 79
Neurotransmitter Receptors and Second-Messenger
Systems 98
There are two major families of neurotransmitter
receptors 99
Second messengers work by activating specific
protein kinases within a cell 102
Tyrosine kinase receptors mediate the effects of
neurotrophic factors 104
Pharmacology of Synaptic Transmission 104
Synaptic Plasticity 105
The Endocrine System 107
Endocrine glands can secrete multiple hormones 107
Mechanisms of hormone action vary 109
Why is the endocrine system important to
pharmacologists? 111
Box 3.2 Pharmacology in Action Sex Hormones and
Drug Abuse 112
Multiple Neurobiological Techniques for Assessing
the CNS 134
Stereotaxic surgery is needed for accurate in vivo
measures of brain function 134
Neurotransmitters, receptors, and other proteins can
be quantified and visually located in the CNS 138
New tools are used for imaging the structure and
function of the brain 145
Genetic engineering helps neuroscientists to ask
and answer new questions 150
Box 4.3 Pharmacology in Action Transgenic Model
of Huntington’s Disease 153
Behavioral and neuropharmacological methods
complement one another 155

5 Catecholamines 159
Catecholamine Synthesis, Release, and
Inactivation 160
Tyrosine hydroxylase catalyzes the rate-limiting step
in catecholamine synthesis 160
Catecholamines are stored in and released from
synaptic vesicles 161
Catecholamine inactivation occurs through the
combination of reuptake and metabolism 164
Organization and Function of the
Dopaminergic System 166
Two important dopaminergic cell groups are found
in the midbrain 166
Ascending dopamine pathways have been implicated
in several important behavioral functions 167
Box 5.1 Clinical Applications Mutations That Affect
Dopamine Neurotransmission 168
6 Serotonin 189
Serotonin Synthesis, Release, and Inactivation 190
Serotonin synthesis is regulated by enzymatic activity and
precursor availability 190
Similar processes regulate storage, release, and
inactivation of serotonin and the catecholamines 192
Box 6.1 History of Psychopharmacology “Ecstasy”—
Harmless Feel-Good Drug, Dangerous Neurotoxin, or
Miracle Medication? 193
Organization and Function of the
Serotonergic System 196
7 Acetylcholine 213
Acetylcholine Synthesis, Release, and
Inactivation 214
Acetylcholine synthesis is catalyzed by the enzyme
choline acetyltransferase 214
Many different drugs and toxins can alter acetylcholine
storage and release 214
Acetylcholinesterase is responsible for acetylcholine
breakdown 215
Box 7.1 Pharmacology In Action Botulinum Toxin—
Deadly Poison, Therapeutic Remedy, and Cosmetic Aid 216
Contents ix
There are five main subtypes of dopamine receptors
organized into D1- and D2-like families 171
Dopamine receptor agonists and antagonists affect
locomotor activity and other behavioral functions 172
Box 5.2 The Cutting Edge Using Molecular Genetics to
Study the Dopaminergic System 174
Organization and Function of the
Noradrenergic System 177
Norepinephrine is an important transmitter in both
the central and peripheral nervous systems 177
Norepinephrine and epinephrine act through α- and
β-adrenergic receptors 178
The central noradrenergic system plays a significant
role in arousal, cognition, and the consolidation of
emotional memories 179
Several medications work by stimulating or inhibiting
peripheral adrenergic receptors 183
The serotonergic system originates in the brainstem
and projects to all forebrain areas 196
The firing of dorsal raphe serotonergic neurons varies
with behavioral state and in response to rewards and
punishments 197
There is a large family of serotonin receptors, most of which
are metabotropic 198
Multiple approaches have identified several behavioral and
physiological functions of serotonin 200
Box 6.2 The Cutting Edge Serotonin and
Aggression 203
Organization and Function of the Cholinergic
System 219
Cholinergic neurons play a key role in the functioning of
both the peripheral and central nervous systems 219
Box 7.2 The Cutting Edge Acetylcholine and Cognitive
Function 220
There are two acetylcholine receptor subtypes: nicotinic and
muscarinic 222
x Contents
8 Glutamate and GABA 231
Glutamate 232
Glutamate Synthesis, Release,
and Inactivation 232
Neurons generate glutamate from the precursor
glutamine 232
Glutamate packaging into vesicles and uptake after release
are mediated by multiple transport systems 232
Organization and Function of the
Glutamatergic System 235
Glutamate is the neurotransmitter used in many excitatory
pathways in the brain 235
Both ionotropic and metabotropic receptors mediate the
synaptic effects of glutamate 236
Box 8.1 Clinical Applications Fragile X Syndrome
and Metabotropic Glutamate Receptor Antagonists:
A Contemporary Saga of Translational Medicine 239
AMPA and NMDA receptors play a key role in learning and
memory 239
High levels of glutamate can be toxic to nerve cells 245
9 Drug Abuse and Addiction 265
Introduction to Drug Abuse and Addiction 266
Drugs of abuse are widely consumed in our society 266
Drug use in our society has increased and has become more
heavily regulated over time 267
Features of Drug Abuse and Addiction 270
Drug addiction is considered to be a chronic, relapsing
behavioral disorder 270
There are two types of progression in drug use 272
Box 9.1 Of Special Interest Should the Term Addiction
Be Applied to Compulsive Behavioral Disorders That Don’t
Involve Substance Use? 273
Which drugs are the most addictive? 275
Factors That Influence the Development
and Maintenance of Drug Abuse and
Addiction 276
The addiction potential of a substance is influenced by
its route of administration 277
Most abused drugs exert rewarding and reinforcing
effects 277
GABA 251
GABA Synthesis, Release, and Inactivation 251
GABA is synthesized by the enzyme glutamic acid
decarboxylase 251
GABA packaging into vesicles and uptake after release
are mediated by specific transporter proteins 251
GABA is coreleased with several other classical
neurotransmitters 253
Organization and Function of the
GABAergic System 254
Some GABAergic neurons are interneurons, while others are
projection neurons 254
The actions of GABA are primarily mediated by ionotropic
GABAA receptors 254
Box 8.2 Clinical Applications GABA and Epilepsy 255
GABA also signals using metabotropic GABAB
receptors 260
Drug dependence leads to withdrawal symptoms when
abstinence is attempted 280
Discriminative stimulus effects contribute to drug-seeking
behavior 282
Genetic factors contribute to the risk for addiction 283
Psychosocial variables also contribute to addiction risk 285
The factors contributing to drug addiction can be
combined into a biopsychosocial model 287
The Neurobiology of Drug Addiction 289
Drug reward and incentive salience drive the binge–
intoxication stage of drug use 289
The withdrawal/negative affect stage is characterized
by stress and by the recruitment of an antireward
circuit 292
The preoccupation/anticipation stage involves
dysregulation of prefrontal cortical function and
corticostriatal circuitry 294
Molecular neuroadaptations play a key role in the
transition to an addicted state 296
Is addiction a disease? 299

10 Alcohol 307
Psychopharmacology of Alcohol 308
Alcohol has a long history of use 308
What is an alcohol and where does it come from? 309
The pharmacokinetics of alcohol determines its
bioavailability 310
Chronic alcohol use leads to both tolerance and
physical dependence 313
Alcohol affects many organ systems 315
Box 10.1 Pharmacology in Action The Role of
Expectation in Alcohol-Enhanced Human Sexual
Response 315
11 The Opioids 351
Narcotic Analgesics 352
The opium poppy has a long history of use 352
Minor differences in molecular structure determine
behavioral effects 353
Bioavailability predicts both physiological and behavioral
effects 354
Opioids have their most important effects on the CNS and
on the gastrointestinal tract 354
Box 11.1 Clinical Applications Saving a Life: Naloxone
for Opioid Overdoses 355
Opioid Receptors and Endogenous
Neuropeptides 357
Receptor binding studies identified and localized
opioid receptors 357
Four opioid receptor subtypes exist 358
Several families of naturally occurring opioid peptides bind
to these receptors 360
Box 11.2 The Cutting Edge Science in Action 361
Opioid receptor–mediated cellular changes are
inhibitory 365
Opioids and Pain 366
The two components of pain have distinct features 367
Opioids inhibit pain transmission at spinal and
supraspinal levels 369
Other forms of pain control depend on opioids 371
Contents xi
Neurochemical Effects of Alcohol 325
Animal models are vital for alcohol research 325
Alcohol acts on multiple neurotransmitters 327
Alcohol Use Disorder (AUD) 335
Defining alcohol use disorder and estimating its incidence
have proved difficult 335
The causes of alcohol use disorder are multimodal 338
Multiple treatment options provide hope for
rehabilitation 342
Opioid Reinforcement, Tolerance,
and Dependence 374
Animal testing shows significant reinforcing
properties 376
Dopaminergic and nondopaminergic components
contribute to opioid reinforcement 376
Long-term opioid use produces tolerance, sensitization,
and dependence 377
Box 11.3 Of Special Interest The Opioid
Epidemic 379
Several brain areas contribute to the opioid abstinence
syndrome 381
Neurobiological adaptation and rebound constitute
tolerance and withdrawal 381
Environmental cues have a role in tolerance, drug abuse,
and relapse 382
Treatment Programs for Opioid Use Disorder 383
Detoxification is the first step in the therapeutic
process 383
Treatment goals and programs rely on pharmacological sup-
port and counseling 384
xii Contents

12 Psychomotor Stimulants: Cocaine, Amphetamine, and Related Drugs 391

Cocaine 392 Repeated or high-dose cocaine use can produce serious

health consequences 409
Background and History 392 Pharmacological, behavioral, and psychosocial methods are
Basic Pharmacology of Cocaine 393 used to treat cocaine abuse and dependence 409

Mechanisms of Cocaine Action 395 The Amphetamines 412

Acute Behavioral and Physiological Effects Background and History 412
of Cocaine 397 Basic Pharmacology of the Amphetamines 413
Cocaine stimulates mood and behavior 398
Mechanisms of Amphetamine and
Cocaine’s physiological effects are mediated by the Methamphetamine Action 414
sympathetic nervous system 399
Dopamine is important for many effects of cocaine and Behavioral and Neural Effects of
other psychostimulants 399 Amphetamines 415
Brain imaging has revealed the neural mechanisms of Amphetamine and methamphetamine have
psychostimulant action in humans 401 therapeutic uses 415
Several DA receptor subtypes mediate the functional High doses or chronic use of amphetamines
effects of psychostimulants 402 can cause a variety of adverse effects 415
Cocaine Abuse and the Effects of Chronic Cocaine Methylphenidate, Modafinil, and
Exposure 403 Synthetic Cathinones 418
Experimental cocaine use may escalate over time to a
Methylphenidate 418
pattern of cocaine abuse and dependence 403
Chronic cocaine exposure leads to significant behavioral Box 12.2 Clinical Applications Psychostimulants
and neurobiological changes 405 and ADHD 420
Box 12.1 The Cutting Edge Neurochemical Mechanisms Modafinil 422
of Cocaine Tolerance and Sensitization 406
Synthetic Cathinones 423

13 Nicotine and Caffeine 429

Nicotine 430 Nicotine exerts both reinforcing and aversive effects 436
Nicotine produces a wide range of physiological
Background and History 430 effects 438
Basic Pharmacology of Nicotine and Its Relationship Nicotine is a toxic substance that can be fatal at high
to Smoking 431 doses 439
Features of tobacco smoking and nicotine Chronic exposure to nicotine induces tolerance and
pharmacokinetics 431 dependence 440
Features of e-cigarette vaping and nicotine Cigarette Smoking and Vaping 443
pharmacokinetics 431
What percentage of the population are current users of
Nicotine metabolism 432 tobacco and/or e-cigarettes? 443
Mechanisms of Action 432 Nicotine users progress through a series of stages in their
pattern and frequency of use 444
Behavioral and Physiological Effects 433 Box 13.1 The Cutting Edge How Safe Are
Nicotine elicits different mood changes in smokers E-cigarettes? 446
compared with nonsmokers 434 Why do smokers smoke and vapers vape? 447
Nicotine enhances cognitive function 434 Smoking is a major health hazard and a cause of
premature death 450
 Contents xiii

Behavioral and pharmacological strategies are Acute subjective and behavioral effects of caffeine
used to treat tobacco dependence 451 depend on dose and prior exposure 456

Caffeine 455 Caffeine consumption can enhance sports

performance 457
Background 455 Regular caffeine use leads to tolerance and
dependence 458
Basic Pharmacology of Caffeine 456
Caffeine and caffeine-containing beverages pose
Behavioral and Physiological Effects 456 health risks but also exert therapeutic benefits 459
Mechanisms of Action 460

14 Marijuana and the Cannabinoids 467

Background and History of Cannabis Acute Behavioral and Physiological Effects

and Marijuana 468 of Cannabinoids 480
Forms of cannabis and their chemical constituents 468 Cannabis consumption produces a dose-dependent
History of cannabis 468 state of intoxication 480
Marijuana use can lead to deficits in memory and
Basic Pharmacology of Marijuana 470 other cognitive processes 481
THC 470 Rewarding and reinforcing effects of cannabinoids
Cannabidiol 471 have been studied in both humans and animals 482

Mechanisms of Action 472
Cannabinoid effects are mediated by cannabinoid
receptors 472
Pharmacological and genetic studies reveal the
functional roles of cannabinoid receptors 473
Endocannabinoids are cannabinoid receptor agonists
synthesized by the body 474
Cannabis Abuse and the Effects of Chronic Cannabis
Exposure 484
Chronic use of cannabis can lead to the development
of a cannabis use disorder 485
Chronic cannabis use can lead to adverse behavioral,
neurobiological, and health effects 489
Box 14.1 Of Special Interest Beyond Cannabis:
The Rise of Synthetic Cannabinoids 495

15 Hallucinogens, PCP, and Ketamine 501

Hallucinogenic Drugs 502 Pharmacology of Hallucinogenic Drugs 507

Mescaline 502 Different hallucinogenic drugs vary in potency
and in their time course of action 507
Psilocybin 502 Hallucinogens produce a complex set of psychological
and physiological responses 508
Dimethyltryptamine and Related
Tryptamines 504 Most hallucinogenic drugs share a common
indoleamine or phenethylamine structure 509
LSD 504 Indoleamine and phenethylamine hallucinogens
Box 15.1 History of Pharmacology are 5-HT2A receptor agonists 510
The Discovery of LSD 505 Salvinorin A is a κ-opioid receptor agonist 511
NBOMes 506 The neural mechanisms underlying hallucinogenesis
are not yet fully understood 511
Salvinorin A 507 Hallucinogenic drug use leads to adverse effects
in some users 511
Can hallucinogenic drugs be used therapeutically? 513
xiv Contents
PCP and Ketamine 515
Background and History 515
Pharmacology of PCP and Ketamine 516
PCP and ketamine produce a state of dissociation 516
PCP and ketamine are noncompetitive antagonists of
NMDA receptors 517
16 Inhalants, GHB, and Anabolic–Androgenic Steroids 527
Inhalants 528
Background 528
Inhalants comprise a range of substances including
volatile solvents, fuels, halogenated hydrocarbons,
anesthetics, and nitrites 528
Abused inhalants are rapidly absorbed and readily
enter the brain 528
These substances are particularly favored by children and
adolescents 529
Behavioral and Neural Effects 530
Many inhalant effects are similar to alcohol
intoxication 530
Chronic inhalant use can lead to tolerance and
dependence 530
Rewarding and reinforcing effects have been
demonstrated in animals 530
Inhalants have complex effects on central nervous
system (CNS) function and behavioral activity 531
Health risks have been associated with inhalant abuse 532
Gamma-Hydroxybutyrate 534
Background 534
Behavioral and Neural Effects 535
GHB produces behavioral sedation, intoxication,
and learning deficits 535
GHB and its precursors have reinforcing properties 535
Effects of GHB are mediated by multiple mechanisms 536
PCP and ketamine have significant abuse potential 517
Use of PCP, ketamine, or related drugs can cause a
variety of adverse consequences 519
Box 15.2 Pharmacology In Action Getting High on
Cough Syrup 520
Novel therapeutic applications have been proposed for
ketamine 522
Medical and Recreational Uses of GHB 538
GHB is used therapeutically for the treatment of
narcolepsy and alcoholism 538
GHB has significant abuse potential when used
recreationally 539
Anabolic–Androgenic Steroids 540
Background and History 541
Anabolic–androgenic steroids are structurally related to
testosterone 541
Anabolic–androgenic steroids were developed to help build
muscle mass and enhance athletic performance 542
Anabolic–androgenic steroids are currently taken by many
adolescent and adult men 543
Anabolic–androgenic steroids are taken in specific patterns
and combinations 543
Pharmacology of Anabolic–Androgenic
Steroids 545
Research is beginning to unravel the mechanism of action
of anabolic–androgenic steroids on muscle 545
Many adverse side effects are associated with anabolic–
androgenic steroid use 546
Regular anabolic–androgenic steroid use causes
dependence in some individuals 548
Box 16.1 Of Special Interest Anabolic–Androgenic
Steroids and “Roid Rage” 549
Testosterone has an important role in treating
hypogonadism 552

17 Disorders of Anxiety and Impulsivity and the Drugs Used to
Treat These Disorders 559
Neurobiology of Anxiety 560
What is anxiety? 560
The amygdala is important to emotion-processing
circuits 561
Multiple neurotransmitters mediate anxiety 564
Box 17.1 The Cutting Edge Neural Mechanism
Responsible for High Tonic Cell Firing Mediating
Anxiety 566
Genes and environment interact to determine the tendency
to express anxiety 574
The effects of early stress are dependent on timing 576
The effects of early stress vary with gender 577
Characteristics of Anxiety Disorders 579
18 Affective Disorders: Antidepressants and Mood Stabilizers 601
Characteristics of Affective Disorders 602
Major depression damages the quality of life 602
In bipolar disorder moods alternate from mania to
depression 602
Risk factors for mood disorders are biological and
environmental 604
Animal Models of Affective Disorders 608
Models of bipolar disorder 608
Neurochemical Basis of Mood Disorders 610
Serotonin dysfunction contributes to mood disorders 611
Norepinephrine activity is altered by antidepressants 614
Norepinephrine and serotonin modulate one another 615
19 Schizophrenia: Antipsychotic Drugs 633
Characteristics of Schizophrenia 634
There is no defining cluster of schizophrenic
symptoms 634
Etiology of Schizophrenia 636
Abnormalities of brain structure and function occur in
individuals with schizophrenia 636
Genetic, environmental, and developmental factors
interact 639
Contents xv
Drugs for Treating Anxiety, OCD, and PTSD 587
Barbiturates are the oldest sedative–hypnotics 588
Benzodiazepines are highly effective for anxiety
reduction 590
Second-generation anxiolytics produce distinctive
clinical effects 595
Antidepressants relieve anxiety and depression 596
Many novel approaches to treating anxiety are being
developed 597
Neurobiological Models of Depression 615
Box 18.1 The Cutting Edge Epigenetic
Modifications in Psychopathology and Treatment 618
Therapies for Affective Disorders 621
Monoamine oxidase inhibitors are the oldest
antidepressant drugs 621
Tricyclic antidepressants block the reuptake of
norepinephrine and serotonin 623
Second-generation antidepressants have different
side effects 624
Third-generation antidepressants have distinctive
mechanisms of action 625
Drugs for treating bipolar disorder stabilize the highs
and the lows 628
Box 19.1 The Cutting Edge Epigenetic Modifications
and Risk for Schizophrenia 642
Preclinical Models of Schizophrenia 646
Box 19.2 Pharmacology In Action
The Prenatal Inflammation Model of Schizophrenia 648
Neurochemical Models of Schizophrenia 650
Abnormal dopamine function contributes to
schizophrenic symptoms 650
xvi Contents

The neurodevelopmental model integrates anatomical and Dopamine receptor antagonism is responsible for
neurochemical evidence 651 antipsychotic action 655
Glutamate and other neurotransmitters contribute to Side effects are directly related to neurochemical
symptoms 652 action 657
Classic Neuroleptics and Atypical Atypical antipsychotics are distinctive in several ways 660
Antipsychotics 650 Practical clinical trials help clinicians make decisions
about drugs 663
Phenothiazines and butyrophenones are classic
neuroleptics 654 There are renewed efforts to treat the cognitive
symptoms 664

20 Neurodegenerative Diseases 671

Parkinson’s Disease and Alzheimer’s Box 20.1 The Cutting Edge Alzheimer’s Disease:
Disease 672 It’s all in your gut??? 684

Parkinson’s Disease 672 Other Major Neurodegenerative

Diseases 685
The clinical features of PD are primarily motor
related 672 Huntington’s Disease 685
Patients with Parkinson’s may also develop dementia 673 Symptoms 686
The primary pathology of PD is a loss of dopaminergic Only symptomatic treatments are available for HD;
neurons in the substantia nigra 673 none alter disease progression 686
Animal models of PD have strengths and limitations 676
Amyotrophic Lateral Sclerosis 687
Pharmacological treatments for PD are primarily
symptomatic, not disease altering 676 The symptoms and disease progression in ALS
are devastating 687
There are several unmet needs in PD diagnosis
and treatment 677 The loss of motor neurons in ALS is complicated
and poorly understood 687
Alzheimer’s Disease 678 Two medications exist that are approved for ALS
AD is defined by several pathological cellular treatment 688
disturbances 679
Multiple Sclerosis 688
There are several behavioral, health, and genetic
risk factors for AD 681 The symptoms of MS are variable and unpredictable 689
Alzheimer’s disease cannot be definitively diagnosed Diagnosis 689
until postmortem analysis 682 Causes of MS 690
Several different animal models contribute to our Treatments fall into several categories for MS
understanding of AD 683 and can be very effective 691
Symptomatic treatments are available, and several others Box 20.2 Pharmacology in Action Can We Repair
are under study for slowing disease progression 683 or Replace Myelin? 693

Glossary G-1
References R-1
Author Index AI-1
Subject Index SI-1
Preface
When we wrote the preface to the Second Edition of
Psychopharmacology: Drugs, the Brain, and Behavior, we
were struck by the many exciting developments in the
field and the remarkable rate of progress elucidating
the underlying neurobiological mechanisms of psycho-
active drug action. This has not changed over the 5
years since the publication of that edition. The entire
field of neuroscience, including neuropsychopharma-
cology, continues to be driven by technical advances.
Using optogenetics, neurobiologists can activate or
suppress anatomically and molecularly defined popu-
lations of nerve cells with amazing temporal precision.
Neuropharmacologists can visualize the 3-dimensional
structure of neurotransmitter receptors, enabling syn-
thetic chemists to design novel agonist or antagonist
drugs with much greater selectivity than could have
been possible before. And huge projects like the Human
Connectome Project (www.humanconnectomeproject.
org) are using the most advanced neuroimaging tech-
niques to map the detailed circuitry of the living human
brain. Because of these technical innovations, we con-
tinue to add new information to Chapter 4, on Methods
of Research in Psychopharmacology. Readers may choose
to go through the chapter in its entirety to familiarize
themselves with all of the neuropharmacological and
behavioral methods reviewed, or they may choose to
use the chapter as a reference source when they en-
counter an unfamiliar method in one of the book’s later
chapters.
Development and introduction of new pharma-
ceutical compounds continues as well, although the
emphasis has somewhat shifted away from the large
pharmaceutical companies to a greater reliance on
drug discovery efforts by researchers at universities
and medical centers. Statistics show that development
of new drugs for CNS disorders (e.g., schizophrenia,
depression, and Alzheimer’s disease) costs more than
for other kinds of disorders, and the failure rate is sig-
nificantly higher. These data have caused many of the
large companies to downsize their CNS drug discovery
programs. As you read the chapters on drug addiction,
mental disorders, and neurodegenerative disorders, it
will become apparent that new medications for these
disorders are being introduced at a slower rate than
expected, despite ongoing research that continues to
identify potential new molecular targets for pharma-
cotherapy. For this reason, we must admit that excit-
ing advances in understanding the basic structure and
function of the nervous system have not yet led to sim-
ilar progress in treating, much less “curing,” disorders
of this system. We came to the same conclusion when
writing the preface to the Second Edition, so it’s disap-
pointing that the hoped-for surge in medication devel-
opment failed to occur during the intervening period.
As before, this new edition of the text is complete-
ly updated to incorporate the latest research findings,
methodological advances, and novel drugs of abuse.
Regarding the latter, illicit drug labs in the United
States and abroad are working hard to turn out mas-
sive amounts of recreational drugs, whether already
known compounds such as cocaine or fentanyl, or
novel synthetic compounds that can only be identi-
fied by submitting drug seizures to advanced forensic
laboratories for chemical analysis. The national drug
epidemic involving fentanyl, heroin, and other opioid
compounds is discussed in Chapter 11. New and, in
some cases, highly dangerous stimulant and cannabi-
noid drugs are introduced in Chapters 12, 14, and 15
respectively. Most chapters have new opening vignettes
and breakout boxes, and new photographs, drawings,
and graphs have been added to bring attention both
to updated material and to completely new topic areas
for discussion.
Importantly, in preparing this next edition of the
book we have maintained our conviction that a deep
understanding of the relationship between drugs and
behavior requires basic knowledge of how the nervous
system works and how different types of drugs interact
with nervous system function (i.e., mechanisms of drug
action). We have also continued to present the methods
and findings from behavioral pharmacological studies
using animal models alongside key studies from the
human clinical research literature. Pharmacologists
must depend on in vitro preparations and laboratory
xviii Preface
animal studies for determining mechanisms of drug
action, for screening new compounds for potential ther-
apeutic activity, and, of course, for basic toxicology and
safety testing. In cases in which clinical trials have al-
ready been performed based on promising preclinical
results, both sets of findings are presented. In other
instances in which clinical trials had not yet been un-
dertaken at the time of our writing, we have striven to
point you toward new directions of drug development
so that you can seek out the latest information using
your own research efforts.
A new point of emphasis in the text concerns neural
circuits as mediators of behavior and as targets of drug
action. As implied above in referring to the Human
Connectome Project, focusing on circuits instead of
cells as the nervous system’s functional units is the con-
temporary way to think about how our brains control
our actions, and how drugs, whether recreational or
medicinal, alter our subjective awareness and behavior.
The Third Edition of Psychopharmacology: Drugs, the
Brain, and Behavior retains the same four-section orga-
nization as the previous editions. Chapters 1 through 4
provide extensive foundation materials, including the
basic principles of pharmacology, neurophysiology
and neuroanatomy, cell signaling (primarily synaptic
transmission), and current methods in behavioral as-
sessment and neuropharmacology. An increased use
of clinical examples demonstrates the relevance of the
material to real-life issues. Chapters 5 through 8 de-
scribe key features of major neurotransmitter systems,
including the catecholamines, serotonin, acetylcholine,
glutamate, and GABA. These are the neurotransmitter
systems most often associated with psychoactive drug
effects, and presentation of their neurochemistry, anat-
omy, and function lays the groundwork for the chap-
ters that follow. Chapters 9 through 16 cover theories
and mechanisms of drug addiction and all the major
substances of abuse. Finally, Chapters 17 through 20
consider the neurobiology of neuropsychiatric and neu-
rodegenerative disorders and the drugs used to treat
these disorders. Among the neuropsychiatric disor-
ders, special emphasis is placed on affective disorders
such as major depression and bipolar disorder, various
anxiety disorders, and schizophrenia. Bulleted interim
summaries highlight the key points made in each part
of the chapter. New to this edition, study questions are
provided at the end of each chapter to assist students
in reviewing the most important material. Finally, a
dedicated website for the book (oup-arc.com/access/
meyer-3e) is available that offers Web Boxes (advanced
topics for interested readers), study resources such as
flashcards, web links, and animations that visually
illustrate key neurophysiological and neurochemical
processes important for psychopharmacology.
It has been our privilege in the first two editions
of Psychopharmacology: Drugs, the Brain, and Behavior to
introduce so many students to the study of drugs and
behavior. With this new and updated edition, we hope
to continue this tradition and perhaps inspire some of
you to continue your studies in graduate school and
join the thousands of researchers worldwide who are
working to better understand and ultimately defeat ill-
nesses like addiction, depression, schizophrenia, and
Alzheimer’s disease.

Acknowledgments
This book is the culmination of the efforts of many
dedicated people who contributed their ideas and hard
work to the project. We’d like to thank and acknowl-
edge the outstanding editorial team at Sinauer Asso-
ciates: Sydney Carroll, Martha Lorantos, and Danna
Lockwood, thank you all for your suggestions for
improving the Third Edition, your help and guidance
throughout the process of writing and revising, and not
least for your patience (textbook writing is a slow pro-
cess when one is simultaneously teaching, conducting
research, and meeting administrative responsibilities).
You were unwavering in your vision to produce the
best possible psychopharmacology textbook. Mark Sid-
dall did a superb job of seeking out just the right photo-
graphs for the book. We are indebted to other key staff
members of Sinauer Associates who worked on this
project, including Chris Small, Ann Chiara, and Joan
Gemme. And we must acknowledge Dragonfly Media
Group for the beautiful job rendering the illustrations.
The following reviewers contributed many excel-
lent suggestions for improving the book:
Joel Alexander, Western Oregon University
Sage Andrew, University of Missouri
Susan Barron, University of Kentucky
Ethan Block, University of Pittsburgh
Kirstein Cheryl, University of South Florida
Matt Clasen, American University
Patricia DiCiano, The Centre for Addiction and
Mental Health and Seneca College
Preface xix
Henry Gorman, Austin College
Bill Griesar, Portland State University
Joshua Gulley, University of Illinois at Urbana-
Champaign
Matt Holahan, Carleton University
Phillip Holmes, University of Georgia
Michael Kane, University of Pennsylvania
Thomas Lanthorn, Sam Houston State University
Lauren Liets, University of California, Davis
Ilyssa Loiacono, Queens College
Margaret Martinetti, The College of New Jersey
Janice McMurray, University of Nevada, Las Vegas
M. Foster, Olive Arizona State University
Robert Patrick, Brown University
Anna Rissanen, Memorial University, Newfoundland
Margaret Ruddy, The College of New Jersey
Jeffrey Rudski Muhlenberg College
Lawrence Ryan Oregon State University
Fred Shaffer Truman State University
Evan Zucker Loyola University
Most of all, we are indebted to our spouses, Melinda
Novak and Ray Rosati, who supported and encouraged
us and who willingly sacrificed so much of our time
together during this lengthy project. Linda gives special
thanks to her husband Ray for providing extensive ed-
itorial advice during the final production period.
Media and Supplements to accompany
Psychopharmacology, Third Edition
FOR THE STUDENT
Companion Website
(oup-arc.com/access/meyer-3e)
The Psychopharmacology, Third Edition companion
website provides students with a variety of resources
to help them learn and review the material presented
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charge and includes the following resources:
•• Chapter Outlines & Summaries provide detailed
reviews of each chapter.
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described in the textbook.
•• Web Boxes include coverage of novel and cutting-
edge topics useful for special discussion.
•• Web Links list suggested websites and other
online resources related to each chapter.
•• Suggested Readings provide additional references
and readings for each chapter.
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of new terms introduced in the textbook.
•• An online Glossary provides definitions for all
textbook bolded terms.
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•• Textbook Figures and Tables (JPEG): All of the
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Psychopharmacology
Drugs, the Brain, and Behavior
THIRD EDITION
 CHAPTER 1

Maggot therapy can be used to clean wounds and prevent infection.

(PA Images/Alamy Stock Photo.)
Principles of Pharmacology
WILLIAM S. BAER (1872–1931) WAS AN ORTHOPEDIC SURGEON at Johns
Hopkins University, where he established the orthopedic department and
led it for most of his life, training many of the outstanding orthopedists of
the day. During World War I Baer observed that soldiers who had severe
and deep flesh wounds did not have the fever associated with infection
and showed little of the expected necrotic (dead) tissue damage if there
was a significant presence of maggots (fly larvae) in the wounds. Although
it had been believed that early peoples (Australian aborigines and Mayan
Indian tribes) and others throughout history had used maggots to clean
wounds, it was Baer who once again recognized their importance, espe-
cially in tense battlefield conditions where infection was especially hard to
treat. Apparently the maggots ingested the dying tissue but left healthy
tissue intact. Baer, upon doing further “pharmacological” experiments,
showed that his hospitalized patients with severe and chronic bone infec-
tions showed remarkable recovery after being treated with maggots—the
inflamed and dying tissue was ingested, leaving wounds clean and healthy,
and new tissue formed. As long as the maggots were sterilized, secondary
infections were avoided. After his research, “maggot therapy” became
popular and was used throughout the 1930s and 1940s until penicillin was
established as an easier treatment for infection. However, it has been sug-
gested that in modern times, maggot therapy will be reintroduced to treat
those wounds infected with antibiotic-resistant bacteria. Presently in the
European Union, Japan, and Canada, maggots are considered “medicinal
drugs,” and in 2005 the U.S. Food and Drug Administration approved the
use of maggots as a medical “device.”
What actually causes the amazing healing process is not entirely clear,
but pharmacologists are beginning to understand that maggot secretions
suppress the immune system and reduce inflammation, and they may also
enhance cell growth and increase oxygen concentration in the wound.
This is certainly not the first time pharmacology has returned to earlier
forms of therapeutics, but the science now can isolate and identify those
components that lead to healing. n
4 Chapter 1
Pharmacology:
The Science of Drug Action
Pharmacology is the scientific study of the actions of
drugs and their effects on a living organism. Until the
beginning of the last century, pharmacologists stud-
ied drugs that were almost all naturally occurring sub-
stances. The importance of plants in the lives of ancient
humans is well documented. Writings from as early as
1500 bce describe plant-based medicines used in Egypt
and in India. The Ebers Papyrus describes the prepa-
ration and use of more than 700 remedies for ailments
as varied as crocodile bites, baldness, constipation,
headache, and heart disease. Of course, many of these
treatments included elements of magic and incantation,
but there are also references to some modern drugs
such as castor oil and opium. The Chinese also have a
very long and extensive tradition in the use of herbal
remedies that continues today. World Health Organi-
zation estimates suggest that in modern times, as many
as 80% of the people in developing countries are totally
dependent on herbs or plant-derived medicinals. And
in 1999, in the United States, modern herbal medicines
and drugs based on natural products represented half
of the top 20 drugs on the market (Hollinger, 2008).
Many Americans are enamored with herbal medica-
tions despite limited clinical support for their effective-
ness, because they believe these treatments are more
“natural.” Nevertheless, serious dangers have been
associated with some of them. Web Box 1.1 discusses
the benefits and dangers of herbal remedies.
When placed in historical context, it can be seen
that drug development in the United States is in its in-
fancy. The rapid introduction of many new drugs by the
pharmaceutical industry has forced the development of
several specialized areas of pharmacology. Two of these
areas are of particular interest to us. Neuropharma-
cology is concerned with drug-induced changes in the
functioning of cells in the nervous system, and psycho-
pharmacology emphasizes drug-induced changes in
mood, thinking, and behavior. In combination, the goal
of neuropsychopharmacology is to identify chemi-
cal substances that act on the nervous system to alter
behavior that is disturbed because of injury, disease, or
environmental factors. Additionally, neuropsychophar-
macologists are interested in using chemical agents as
probes to gain an understanding of the neurobiology
of behavior.
When we speak of drug action, we are referring
to the specific molecular changes produced by a drug
when it binds to a particular target site or receptor.
These molecular changes lead to more widespread al-
terations in physiological or psychological functions,
which we consider drug effects. The site of drug ac-
tion may be very different from the site of drug effect.
For example, atropine is a drug used in ophthalmology
to dilate the pupil of the eye before eye examinations.
Atropine has a site of action (the eye muscles of the iris)
that is close to the site of its ultimate effect (widening
the pupil), so it is administered directly to the eye. In
comparison, morphine applied to the eye itself has no
effect. Yet when it is taken internally, the drug’s action
on the brain leads to “pinpoint” pupils. Clearly, for
morphine, the site of effect is far distant from the site
of its initial action.
Keep in mind that because drugs act at a vari-
ety of target sites, they always have multiple effects.
Some may be therapeutic effects, meaning that the
drug–receptor interaction produces desired physical
or behavioral changes. All other effects produced are
referred to as side effects, and they vary in severity
from mildly annoying to distressing and dangerous.
For example, amphetamine-like drugs produce alert-
ness and insomnia, increased heart rate, and decreased
appetite. Drugs in this class reduce the occurrence of
spontaneous sleep episodes characteristic of the disor-
der called narcolepsy, but they produce anorexia (loss
of appetite) as the primary side effect. In contrast, the
same drug may be used as a prescription diet control
in weight-reduction programs. In such cases, insomnia
and hyperactivity are frequently disturbing side effects.
Thus therapeutic and side effects can change, depend-
ing on the desired outcome.
It is important to keep in mind that there are no
“good” or “bad” drugs, because all drugs are just chem-
icals. It is the way a drug is procured and used that de-
termines its character. Society tends to think of “good”
drugs as those purchased at a pharmacy and taken at
the appropriate dosage for a particular medicinal pur-
pose, and “bad” drugs as those acquired in an illicit
fashion and taken recreationally to achieve a desired
psychological state. Even with this categorization, the
differences are blurred because many people consider
alcohol to be “bad” even though it is purchased legally.
Morphine and cocaine have legitimate medicinal uses,
making them “good drugs” under some conditions,
although they can, when misused, lead to dangerous
consequences and addiction, making the same drugs
“bad.” Finally, many “good” prescription drugs are ac-
quired illicitly or are misused by increasing the dose,
prolonging use, or sharing the drug with other indi-
viduals, leading to “bad” outcomes. As you will read
in later chapters, the ideas of Americans about appro-
priate drug use have changed dramatically over time
(see the sections on the history of the use of narcotics
in Chapter 11 and cocaine in Chapter 12).
Many of the drug effects we have described so far
have been specific drug effects , defined as those
based on the physical and biochemical interactions of
a drug with a target site in living tissue. In contrast,
nonspecific drug effects are those that are based not
on the chemical activity of a drug–receptor interaction,
Another random document with
no related content on Scribd:
across the clouds like spears of fire threatening revenge. The stone
has remained to the present day, and is known by the name of
“Koreno kardjo (dog) gambi” (stone).

PLATE XXXIV

The “Tjilbakuta” of the great emu ceremony, Arunndta tribe.

 “The moment the sacred object is completed, the Illiyakuta delegates
one of his group to act as its attendant or guardian.”

The snake is an important character in the mythology of practically

every tribe of Australia; in fact most of the permanent water-holes
are supposed to be inhabited by great serpents which guard the
supplies, destroy unlawful consumers or polluters thereof, and
frequently communicate with those spirit ancestors of the tribes who
are descended from the original snake-man still living in the sky. In
many cases the mythic snakes can be recognized in some
characteristic features of the landscape. Take, for instance, the great
artesian spring near Coward Springs Station which is known as
Blanche Cup. This is looked upon as the mouth of a snake, while the
hill immediately at the back of it (Mt. Hamilton) is its head. In
consequence the formation is called “Worma-Kadiabba” (snake’s
head) by the local Arrabonna tribe. The natives have a dread of
these imaginary snake-monsters and prefer not to visit a water-hole
at night; in fact, at any time, day or night, they feel safer in the
company of a man who is “related” to the snake, because he can
protect them and give them the right of approach. The snake is
possessed of evil and will molest any but its totemic “relatives.”
The fundamental conception of the kobong (or totem), so far as
the Australian aboriginal is concerned, is of a religious nature. In the
beginning of all things, the Aluridja say a number of exalted
creatures of human form came out of the earth and were gracious to
their tribes-people. Then appeared a menace in the shape of a
gigantic dog which chased the good people from one place to
another, until they decided to adopt the forms of various animals and
plants, and thereby became either too fleet for the dog or were not
recognized by it. Other good people now descended from the hills
and drove the dog back to its hiding place in a cave where the evil
spirit dwells. The newcomers kindled a fire at the mouth of the cave
and kept the evil beings in captivity whilst the original Deities re-
assumed the human form. Ever after, however, these good creatures
were able to alter their appearance from human to animal at will; but
each individual in his choice adhered to the particular animal or plant
which had saved him from the ravages of the great evil dog.
Eventually they formed themselves into flat slabs of stone or wood,
upon the surfaces of which they scratched the emblems of their
animal representation and the traditions of their long wanderings on
earth. The spirits of these Deities now live in the sky but can return
at any time to re-enter the slab generally known as the “tjuringa.”
Among the Minning at Eucla the larger of these objects are known as
“wagal-wagal,” the smaller as “bobi,” whilst further west, in the
Laverton district, “kaidi” is the prevailing word. It is true, the tjuringa
is not known to all tribes; in which case the Deities are supposed to
have entered such natural objects as rocks, hills, and conspicuous
trees.
The Roper River natives believe that their deified forbears were
molested not by a dog, but by a hideous old woman or witch, who,
by the influence of evil, entrapped them and subsequently ate them.
On one occasion, however, a party of warriors were successful in
decoying her away from her haunts and slaying her. The jubilant
victors decided to cut out the old woman’s tongue as a trophy, but as
they were thus engaged, the tongue flew out of the mouth and spun
round in the atmosphere above them, making a terrible noise as it
did so. The men chased the tongue, but it flew towards a beefwood
tree and embedded itself deeply in the butt; in vain they looked for it
and tried to cut it out; it had become part of the tree. Before
returning, however, the men took a piece of wood out of the tree,
shaped like the woman’s tongue, which they tied to a piece of human
hair-string and swung round their heads with joy. Behold their mixed
feelings of delight and fear when the piece of wood began to howl
with a voice like that of the slain witch! The tribe retained that piece
of wood as a sacred memento of their victory, and they gave to it the
name the witch was known by, namely “Kunapippi.” Nowadays this
object is the equivalent of the central Australian tjuringa.
All tribes recognize the existence of deified ancestors, now real or
spiritual, whom they regard as sacred and worship accordingly. All
ancestors stand in a definite, intricate, and intimate relationship to
some animal, plant, water-hole, or other natural object which they
have at some time or other represented; some indeed in the first
place appeared as animals and later took the human form. They are
now looked upon as being those powers who by virtue of sacred
ceremonial can produce the species they have at some time
incarnated, in plenty or allow it to proliferate. As a matter of fact,
some of the sorcerers of the tribes often declare that they can see
the inside of a sacred rock or tjuringa teeming with young, ready to
be produced.
The Arunndta refer to their “Knaninja” (i.e. “totem” Deities) as
“Altjerrajara,” meaning the Supreme Number; the Aluridja as
“Tukurata” or “Tukutita”; and the Dieri as “Muramura.”
Just as the “totem” ancestor is connected with an animal, plant, or
other natural object, and is embodied in the sacred form of the
tjuringa, so the individual who traces his descent from such ancestor
recognizes a close and mysterious affinity between himself and the
tjuringa which has become his by heredity; henceforth it becomes his
sacred talisman which protects him from evil and procures for him
the means of maintaining his existence.
The emblematic representation of the deified ancestor, based
upon the form of an animal or plant living to-day and in some way
“connected” with the individual, is the “kobong” of the north-western
tribes first referred to by Sir George Grey.
The “totem” is very dear and sacred to the native, and is religiously
protected by him. I well remember on one occasion on the Alberga
River I discovered a small black and yellow banded snake which I
killed. An Aluridja man who was attached to the party at the time was
greatly shocked at this, and, with genuine sorrow, told me that I had
killed his “brother.” Turning to an Arunndta he lamented aloud:
“Kornye! Nanni kallye nuka kalla illum,” which literally translated
means: “Oh dear! This brother of mine is dead.”
One thing is always essential and that is that a native performs
frequent, prolonged, and reverential ceremonies, remote from the
women and children, and in the presence of his tjuringa. Under these
conditions the tjuringa is believed to have powers similar to those of
the Deity it embodies.
 When not in use, the tjuringas are stored in caves, the entrances
to which are small and not easily discernible; the ground is
proclaimed taboo to any but initiated tjuringa holders and is strictly
regarded as a sanctum sanctorum. Although the sticks and stones
are the individual property of the tribesmen, the objects are generally
kept together, and only brought out during a religious ceremony. The
old men are the authorized custodians of the sacred collection. The
female tjuringas are included, because even though a woman may
possess one, she must never see it; if she does, accidentally or
otherwise, she is in imminent danger of being killed. No unauthorized
hunter is allowed near the prohibited area under any pretext at all;
even if an animal he has wounded should by accident make for the
sacred ground to breathe its last, the hunter is required by tribal law
and usage to sacrifice it to the divine factors incorporated in the
tjuringa, by leaving it on the spot.
 PLATE XXXV

Flashlight photograph of “Illiya Tjuringa” or great emu ceremony, Arunndta

tribe.

“The chief emu man is distinguished by an extra large head-dress called the
‘Illiya Altjerra Kuta.’...”
 When on the warpath, a warrior always craves to carry his tjuringa
with him, even though this is not always possible. He firmly believes
that with the talisman kept on his person, or at any rate knowing that
it is nearby, no deadly missile thrown by an enemy will penetrate his
body. The mere knowledge of the fact that his opponent has a
tjuringa with him, and he not, is sufficient to make a coward of the
bravest fighter. Should he be wounded or take ill, one of his “totem-
brothers” endeavours to produce a tjuringa, from which, if the
medicine man considers it necessary, a little powder is scraped and
handed to the patient to swallow with water.
With regard to “totem” animals which form the objects of hunting
expeditions, a man is allowed to kill and eat thereof with some
restriction. He must kill only one animal at a time, and only in
accordance with the method prescribed by the tribal fathers and
handed down to them by tradition. This is usually a straightforward
hunting method, with as little loss of blood as possible. If much blood
should flow, the hunter is obliged to cover it without delay with sand.
If possible, other men should cut up the carcase, and only certain
portions be handed to the “brother” of the slain animal.
Each tribe has an endless variety of objects (animal, vegetable,
terrestrial, meteoric, mythic, and so forth), which may figure as a
“totem.” Any one of these may be the primary motive of a separate
cult or sacred ceremony, but here again the variety is usually
reduced to the number corresponding to the most sanctimonious and
most useful creations affecting the affairs of the particular tribe.
The ceremonies take the form of either a direct worship or a
prayer for increased productivity of a certain plant or animal, either
being offered to the Knaninja or “totem” ancestors living as spirits in
the sky. Usually the two ideas are embodied in one grand ceremony,
and the method of procedure is governed by tradition. Such
ceremonies have been particularly elaborated by the Arunndta tribe,
who refer to them by the same name as that of the sacred object,
namely “Tjuringa”; less frequently they call them “Intitjuma,” the latter
name being applied more to ceremonies without worship.
 The Tjuringa ceremonies are divided into grades according to their
importance and sacredness. The water ceremony is ordinarily called
“Kwatje Tjuringa,” but if the “totem” spirit ancestor is invoked to
attend, it goes by the name of “Kwatje Tjuringa Knaninja”; if the
principal spirit ancestor is assumed to be present, the title becomes
“Kwatje Tjuringa Knaninja Knurrendora”; and finally the most sacred
water ceremony of all is the “Kwatje Tjuringa Altjerra Knaninja
Knurrendora.”
As a typical illustration we shall discuss the “Illiya Tjuringa” or
Great Emu Ceremony of the eastern Arunndta groups. The date of
the performance is decided by the senior emu “brother” of the tribe,
the oldest member who claims to be related to the Illiya Knaninja.
Somewhat extensive preparations are made beginning a few days
prior to the opening event. Only fully initiated men take part, but the
women are allowed to witness certain of the most awe-inspiring
stages from a distance. Whilst the younger men are out collecting
leaves, out of which they make the down later to adorn the bodies of
the performers, the older men prepare the sacred ground. Others
slay a number of brown hawks, off which they pull the feathers and
then pluck the down. A suitable site having been selected, the old
men clear it by removing all grass and bush from the surface and
smoothing the sand with their feet. The “brothers” who claim
relationship alike to the great Emu-Man, the Emu-Knaninja, and the
emu itself, thereupon proceed to anoint the sacred ground with their
blood, for which purpose they puncture the median basilic vein of the
forearm with a quartzite chip and allow the fluid of kinship to sprinkle
upon the sand. It is surprising to see the amount of blood sacrificed
by the men on occasions like this; and time after time, when such is
required, the process is repeated. By examining the forearms of an
old stager, one can usually count a number of small scars along the
course of a vein indicating places where a perforation has at different
times been made. A venesection is made after much the same
manner among the various tribes.
The following Arunndta method will serve as an example. A
ligature of hair-string is in the first place tied tightly round the upper
arm, a little above the biceps muscle, after the style of a tourniquet to
check the flow of blood in the veins and thereby distend the vessels.
The man then makes a small longitudinal cut through the skin and
punctures the vein beneath it lengthwise; the blood spurts forth
immediately and is collected in the handle-pit of a shield. When the
flow is to be stopped, the native removes the ligature, and this in
most cases is all that is needed. Should, however, the blood
continue to come, he places a small amount of down over the
incision and presses it against the vein, or winds three or four
strands of fur-string around it. The little pad of down is usually left on
the arm until it dries and falls off. None of the women are allowed to
witness this operation, which is called “Ilgarukna.” The blood, when it
is to be used as an adhesive for the down-decoration, is applied with
a small brush (“ipinja”) made of twigs tied together with fur-string.
Vide Plate XXXIII.
The principal among the emu group is called “Illiyakuta,” and it is
he who directs the performance. He takes his followers to a secluded
place, such as a clump of timber or down a creek-bed, and there the
wooden tjuringas belonging to the ceremony are produced and
painted afresh with red ochre and emu fat.
Down is made out of the white, felty leaves and twigs of Kochia
bush, which the Arunndta call “kemba.” Small quantities of these are
placed upon a flat slab of stone and pounded with a pebble. The
fluffy material which results is next mixed and rubbed by hand with
powdered kaolin or ochre according to the colour required, the white
being known as “wadua,” the red as “wanjerra.”
A sacred object is now constructed which encloses the painted
tjuringas and is called the “Tjilbakuta.” It is about three feet high and
is made in the following way. The tjuringas are laid one on top of the
other and bound together with many lengths of human hair-string,
which completely obscure the shape of the separate pieces. A thick
layer of the stalks of the kangaroo grass (Anthistiria) is laid around
the parcel and kept in position with a few lengths of twine, and then
the whole structure is covered with great masses of human hair-
string wound spirally from top to bottom. A cylinder results which is
decorated with alternate vertical bands of red and white vegetable
down. Into the top of this Tjilbakuta one bundle of emu feathers and
one of black cockatoo tail-feathers are stuck; and often additional
plumes are hung beneath them. The moment the sacred object is
completed, the Illiyakuta delegates one of his group to act as its
attendant or guardian. For the time being his body is decorated with
symmetrically placed, curved ochre bands upon the chest and
vertical bands down the arms; at a later stage he ornaments his
body more elaborately, prior to taking part in the principal
performance; but all the time he remains in his place of hiding beside
the Tjilbakuta. Vide Plate XXXIV.
At the sanctified place close by the other men have been stacking
firewood at different points to illuminate the proceedings during the
evening. Occasionally, too, the Illiyakuta group of men cover a
portion of the ground with a coloured emblem of the traditional emu.
Early in the afternoon of the festive day the men who will take part
in the ceremony at night begin to prepare themselves. Many of the
non-performers help them.
Large quantities of down, both vegetable and birds’, are used to
decorate the bodies. The design is shaped much like a cobbler’s
apron, extending from the neck down the front to the level of the
knees. The greater part of this surface is red, but it is lined with white
and split along the centre by two parallel lines of white. The back is
not decorated at all. The entire surface of the face, including the eye-
lids and beard, is thickly covered with down which is white, except
for an oval red patch around the mouth.
The principal attraction, however, of the sacred emu ceremony is
the head-dress, which is both elaborate and imposing. To prepare it,
the attendant combs back the actor’s hair with his fingers, and
interlaces it with stalks of grass and small twigs in such a way that a
tall conical structure results right on top of the head. This is made
secure and of a uniform exterior by winding much human hair-string
around it, at the same time taking in a plume of emu feathers at the
apex of the cone. The headgear is completely enveloped in red and
white down, extending upwards from the head as alternate vertical
bands. The chief emu-man is distinguished by an extra large
headdress called the “Illiya Altjerra Kuta”; this measures a good
three feet in length, and it embodies, between the apex and the emu
plume, deeply enshrouded with hair-string and down, the sacred
“Illiya Tjuringa.” Other members who are of the same rank as the
“Tjilbakuta” guardian, wear their insignia beneath the emu feathers in
the form of a sickle-shaped rod, which carries at each of its points a
tuft of white cockatoo feathers. All performers cover their person with
a dog-tail appendage which hangs from a thin waistband of human
hair-string. And lastly, they all tie bundles of eucalyptus twigs, with
the leaves attached, to their legs just above the ankles. If possible,
old or half-dried leaves are selected in order that a more pronounced
rustling is produced when the men move about; the noise is made to
imitate the rustle of the wiry feathers of an emu. Vide Plate XXXV.
At nightfall the Tjilbakuta is removed from the hiding place and
planted on the edge of the ensanguined patch. The guardian is thus
given an opportunity to slip away and to attend to his ceremonial
toilet, which is similar to that of the rest of the Tjilbakuta group. When
he returns, the performance is about to begin, and all except he
leave the ground.
The stacks of wood are set fire to by invisible hands, and, so soon
as the flames flare upwards, the silence is broken by the booming
note of a bull-roarer, which is produced some distance off in the
bush.
The Tjilbakuta guardian sits beside the object like a statue, with
his eyes rivetted to the ground immediately in front of him. From
behind him the thud of stamping feet and the rustle of dry leaves
announce the coming of the official performers, while from the other
side the non-performing members step from the darkness and take
up their position by squatting between two fires. When the decorated
men come into view, the latter start beating their boomerangs
together in perfect time to the stamping of the feet of the advancing
actors. They come as a body of five or six rows, one behind the
other, each man holding his hands locked behind his back and
uttering a deep guttural note resembling a pig’s grunt. The folded
hands held over the stern represent the tail, the guttural noise the
call of the emu.
 The Illiyakuta, wearing the tall Illiya Altjerra Kuta, is in the front
row, and he is attended on either side by a Tjilbakuta man. The chief
now starts a chant: “Immara janki darrai,” and all the others,
including the sitting men, join in; the same is repeated several times.
When the two parties are opposite each other, the performers
quicken the pace of their stamping and extend their arms sideways,
thereby widening their ranks. After this they retreat to behind the
Tjilbakuta and one hears a shrill chirping note resembling the cry of a
young emu.
The interpretation of this act needs no special elucidation. The
decorated performers are those of the tribe’s manhood who, in all
matters pertaining to the emu, have a right to communicate, through
the Tjilbakuta, with the astral emu ancestor living in the great
celestial domain of the ancestral spirits, which is known as
“Altjerringa.” They are invoking the benign Knaninja or originator of
their particular “totem” species to increase the numbers of emu on
earth for the exclusive benefit of their tribe. It is the Illiyakuta who
imagines that he receives the favourable response from above, and,
when it comes, it is he who imitates the cry of a young emu. It often
happens, however, that the chief persuades himself to believe that
the Great Spirit had not heeded the appeal, in which case the last-
mentioned cry is wanting. The ceremony is repeated time after time.
Altjerringa, it will be observed, is a compound word consisting of
“Altjerra,” the Supreme Spirit, and “inga,” a foot or trail. The implied
idea is that Altjerringa is the “walk-about” of the spirit ancestors,
where they walk, and have always walked, and where the spirits of
all tribes-people eventually hope to find their way.
After this act, the performance becomes less restrained and takes
more the form of a corrobboree. Some of the men seize firebrands
from the burning stacks and hurl them in the direction of the
women’s camp. From the moment of the sounding of the bull-roarer
at the beginning of the ceremony until now the women sat huddled
together, with their faces buried in their hands, thoroughly cowed by
the portentous happenings. When the firebrands come whizzing
through the air and crash into the branches of the trees around them,
sending sparks flying in all directions, they are almost beyond
themselves with fear. But just at this juncture the men call upon them
to look towards the festive ground and behold them dancing. In
obedience to the order, the women’s fears are dispelled and soon
superseded by a noticeable enravishment. They feast their eyes
upon the array of manhood in gala dress, and it is not long ere they
pick up the rhythm of a dance by beating time to the step. Provided
the Tjilbakuta has been removed to a place of secrecy, well out of
reach of accidental discovery, the men entreat the women to come
up and join in the song. Thus the sublime is eventually reduced to
commonplace, and the remainder of the night passes in joviality.
To refer briefly to a vegetable ceremony, we shall select the yam
or “Ladjia Tjuringa Knaninja.” The preparations are much the same
as those of the emu ceremony. An enclosure is first made in a
secluded spot with branches, in the centre of which the “totem” or
Knaninja “stick” is erected. Several men immediately set about to
decorate it with vegetable down as previously described. The design
in this case consists of vertical rows of red circles upon a yellow
ochre background. In addition, a large plume of split eagle-hawk
feathers is stuck into the top of the stick. All ordinary performers
wear conical head-gears or “tdela” made of Cassia twigs, into the
apices of which tightly bound bundles of grass stalks (“gortara”) are
fixed carrying plumes of emu feathers (“mangalingala”) (Plate
XXXVI, 1). Other men have squat, cylindrical bark structures called
“elbola” placed over their heads, which are elaborately decorated
with vertical coils of human hair-string and coloured down.
One of the principal actors represents the “Kuta Knaninja.” His
head-gear consists of two long kutturu, tied together with hair-string
and completely covered with gum leaves, the whole being
subsequently besmeared with blood and decorated with coloured
down. As the assistants are dressing this character, they keep up a
chant sounding like “Winni kutcherai.” Vide Plate XXXVI, 2.
The leading figure is the “Ingada Ladjia Knaninja,” who wears a tall
vertical head-piece which contains the tjuringa of the Ladjia
Knaninja. The tjuringa is, however, not visible, but is covered with
pieces of bark, securely tied over it with hair-string, the whole being
richly decorated with vertical bands of red and white down.
 The Great Spirit of the Yam, called “Knaninja Tjilba Ladjia,” when
he leaves Altjerringa, takes up his abode in a cave near Mount
Conway, where the tjuringas are kept, but at night, before the fires
are lit, he is supposed to come to the ceremonial ground and occupy
the decorated “totem” stick described above. During the performance
he is surrounded by all the ordinary performers, who are known as
“Tjilba Ingarrega,” and are directly under the guidance of the Ingada
and Kuta Knaninja.
A group of men who are not decorated sit near one of the fires and
sing while the performers are thus encircling the Ladjia stick:

“Imbanai yinga
Wi ma bana Ladji di bana
Yammana wi ma bana
Jai ra ja ja
Jai ja ja na
Wi ba na na
De a re a ja betja,”

the voices finally fading away to an almost inaudible whisper.

If the Great Spirit, Ladjia Altjerra Knaninja, is gracious, the tap-root
of the yam will be sent deep down into the earth near the Jay River
and from there spread its laterals all over the country to supply the
needs of the tribe.
When some of the most sacred ceremonies are performed, the
oldest “relatives” of the presiding Knaninja often construct a coloured
drawing upon the consecrated ground, whose purpose is similar to
that of the “totem” stick above described. The drawing is executed in
coloured down, both vegetable and bird. A space of suitable size,
often measuring many feet in length, is cleared of grass and stones,
and sprinkled with water, when it is ready to receive the down. In the
case of, say, the “Ladjia Tjuringa,” the design takes the form of a
number of concentric circles alternately red and white, from the
outermost of which six equally spaced groups of red and white lines
stand out radially. The enclosing border of the design consists
entirely of white down. Vide Plate XXXVII.
 Once constructed, this drawing, which is known as “Etominja,” is
zealously guarded by one of the old men. If, peradventure, an
unauthorized person happens upon the sanctified place, he is killed
and buried immediately beneath the spot occupied by the design;
thereupon the ground is smoothed again and the Etominja re-
constructed. Nobody in camp ever hears what became of the
person, and should any relative track him in the direction of the area
known to be tabooed, he is horror-stricken and runs away.
While the old men are re-constructing the Etominja, they sing to
the Knaninja as follows:

“Yedimidimi
Dakarai pa ma taka,
Pa mitu min jai, jin tarai,
Ja ra nai malgada, ja ranai.”

The next, and probably the most important, group of religious

ceremonies is that dealing with Sex-Worship. For years past
peculiarly shaped stones have been found in caves and among the
possessions of the Australian aborigines whose shape was strikingly
suggestive of a phallus, but hitherto no actual phallic ceremonies
have been observed. It was my good fortune to witness such among
the Aluridja, Arunndta, Dieri, and Cambridge Gulf tribes. From
enquiries made of the old men, it appears that in former days this
form of worship was practised considerably more than it is
nowadays. New stone phallus are rarely made by the present tribes;
those in their possession have generally been inherited from
previous generations. The old men have the phallus in their keeping,
and they are very loth to either produce or part with them.
The natives of the King Sound district in the north-west believe the
origin of the phallus to be as follows: In the early times a scourge
was raging among their forefathers, from the effects of which many
were daily dying, when a hairy man and his mate, a woman of
ordinary human form, came to earth from above. The evil was due to
the exhalation of poisonous breath from the gaping jaws of a green
monster resembling a crocodile. The stranger relieved the sufferers
from the awful curse by showing them how to perform an operation
upon their person which taught them to endure pain and protected
them against future ravages of the pestilence. This great and
benevolent stranger then took his departure, but left his name to
designate the surgical operation which to the present day is
performed upon the male members of the tribe; the name, strange
though it may seem, is “Elaija”; and it is known, at any rate, as far
east as Port George IV. But the tribe had become so weak through
the terrible havoc the disease had wrought that the old men called
him back and entreated him to stay. Elaija, however, took from a
dillybag his female companion was carrying, a stone carved after the
shape of a mutilated member, which he gave the name of
“Kadabba.” When the old men gazed upon this object, they took
fright and appealed to Elaija, but the good fellow had vanished. The
stone has remained with the tribe ever since, and through the divine
property Elaija endowed it with, their threatened extinction was
eluded. Moreover, they continued to practise the operation on all
young men because it made their members like the Kadabba of
Elaija, which they knew had the power of multiplying their kind. And
so the Kadabba became a sacred object whose procreative power
they have learned to worship, thinking that by such observance they
would augment their own capacities of sex. Vide Fig. 7.
 Fig. 7. Stone phallus, Northern Kimberleys, Western Australia (× 1/2).

One often reads, and I was under the same impression myself
until I became better acquainted with the tribes, that the Australian
natives do not connect the knowledge of conception with any
intercourse which might have taken place between the sexes. This I
find is not altogether correct, although usually the younger people
are kept in complete ignorance on the subject. No doubt strangers
are treated similarly when they put any pertinent questions to the old
men on matters of sex. The old men believe in the duality of human
creation, the spiritual and the material; sexuality is regarded as the
stimulus of corporeal reproduction, but the spirit quantity is derived
through mystic and abstract influences controlled by a “totem”-spirit
or Knaninja. Under these circumstances, it is not surprising to note
that the ceremonies of the phallus are transacted principally by the
old men of the tribe who aim at the rejuvenation of their waning
powers.
It is interesting to see the old men preparing for a ceremony which
is to be dedicated to a Knaninja or Spirit of Sex, because they all
endeavour to conceal the white hairs of their beards by rubbing
powdered charcoal into them. The bark of the cork tree (Hakea) is
used for the purpose; pieces of it are charred, crushed between the
palms, and applied where needed. It is astounding what a difference
this process makes to the appearance, and some of the old grey-
beards really look as though they had been made twenty years
younger by magic.
In the eastern MacDonnell Ranges stands a cylindro-conical
monolith whose origin is believed to be as follows: Many generations
ago, the paternal ancestors of the Arunndta walked from a district
situated, as near as one can gather, somewhere in the
neighbourhood of Ediowie; they were known as the “Kukadja,” and
were characterized by the enormous dimensions of their organs.
These old men or Tjilba of the tribe migrated northwards to beyond
Tennant’s Creek and settled in the productive “Allaia” country which
surrounds the Victoria River. In that same district one finds, even at
the present day, cave drawings of human beings with the anatomical
peculiarities referred to (Fig. 8). At a later time, the head-man of the
Kukadja, named “Knurriga Tjilba,” returned southwards to the
Macdonnell Ranges. While roaming the hills, he espied two young
women sitting on the side of a quartzite cliff, and without deliberation
began to approach them. He was in the act of making lewd overtures
when the guardian of the girls, a crow ancestor, caught sight of him
and hurled a boomerang at him. The missile struck the great man
and cut off the prominent portion of his body, which in falling stuck
erect in the ground. The force of the impact was so great that the
man bounced off the earth and fell somewhere near Barrow’s Creek.
He bled so profusely that a clay-pan soon filled with his blood. Thus
his followers found him, and overcome with sorrow they opened the
veins of their arms to mix their blood with his. Then all the members
of the party jumped into the pool and disappeared for ever.
 Fig. 8. Ochre drawing of “Kukadja” men, north of Wickham River, Northern
Territory (× 1/3).

The severed portion of the old man’s body, however, remained just
where it fell and turned to stone. It has long been known as “Knurriga
Tjilba Purra.”
The two young women can also still be detected in the cliff as
prominent rock formations.
The stone has been protected by the tribe as long as the old men
can remember, because they realize that it contains an inexhaustible
number of unborn tribes-people. These mythic, foetal elements are
generally recognized to exist in certain objects of phallic significance,
and are called “rattappa.” The medicine men maintain that they can
at times see the dormant living matter in the stone. It is on that
account that it is regarded as sacred, and every now and then very
secret and worshipful ceremonies are transacted near its base, the
main objects of which are to multiply the future membership of the
tribe and to preserve the sexual powers of the old men.
 The Tjilba Purra naturally figures prominently in some of their
ceremonies. In fact, it is reproduced and worn upon the head of the
leading man during the functions. The sacred effigy consists of an
upright column, about two feet high, composed of a stout bundle of
grass stalks, in the centre of which the tjuringa is contained. It is
decorated with alternating bands of red and white down throughout
its length. This upright column represents the “Tjilba” or revered
ancestor whose spirit is invoked to “sit” in the tjuringa; at the top of it
a plume of wiry emu feathers, well powdered with charcoal (“unjia”)
to give it a youthful appearance, takes the place of the forbear’s hair
and beard. Standing at an angle with the central column, a similar
though slightly smaller structure is intended for the “Purra” or
phallus; it carries a plume of white cockatoo feathers at its end to
represent the glans. Vide Plate XXXVIII, 2.
A landmark, of similar significance as the Tjilba Purra of the
Arunndta, exists on the Roper River in the Northern Territory; it is a
pillar of sandstone known as “Waraka.” Waraka is also the name of
the great Spirit Father of the tribe. In very early times this man came
to earth in a semi-human form, and made the country abound in
game, animals, birds, and fish. Then he found a woman on the
shores of Carpentaria Gulf who remained with him as his wife. Many
children came of the union; and Waraka’s mate has since been
looked upon as the mother of the tribe. The woman’s name was
“Imboromba,” and to this day the tribe takes its name after her.
Warraka had an enormous sex characteristic which was so
ponderous that he was obliged to carry it over one of his shoulders.
Eventually the organ became so huge that Warraka collapsed and
sank into the earth. His burden remained, but turned to stone, and is
now looked upon by the local natives as the great symbol of Nature’s
generative power which first produced their game supplies and then
the original children of the tribe; it is revered accordingly.
The Kukata have a somewhat similar legend of the origin of a
stone of phallic significance, the name of the possessor of the large
organ being “Kalunuinti.”
In the extreme north-western corner of Australia, in the Glenelg
River district, the natural stone is replaced by an artificially