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 vii
Contents  

Extinction 196
Differential Reinforcement 198
Chapter Summary 204
Application Exercises 205
References 206

Chapter 9. Establishing the Evidence Base: Single-Case 207

Experimental Designs
What Is Science? 208
Formulating a Research Question 209
Scientific Experimentation Versus Routine Clinical Work 211
Characteristics of Single-Case Research Designs 214
Types of Single-Case Research Designs 218
External Validity in Single-Case Research 231
Chapter Summary 231
Application Exercises 232
References 233

Chapter 10. Ethics in Scientific Research and Clinical Practice 235

A Brief History of Ethical Issues in Scientific Research 236
The National Research Act of 1974 and the Belmont Report 241
The Distinction Between Scientific Research and Clinical Practice 247
Ethical Issues in Conducting Treatment 248
Chapter Summary 260
Application Exercises 261
References 262

Glossary 265
Index 275
 Preface
Welcome to Behavioral Principles in Commu-
nicative Disorders: Applications to Assessment
and Treatment. This book is an uncommon
text in that it bridges two disciplines in one
practical, application-focused volume.
This interdisciplinary feature has shaped
a valuable and unique contribution to the
current available selection of textbooks for
training in communicative disorders and
in the field of applied behavior analysis
(ABA). The authors of this book have
presented information that is compre-
hensive while remaining focused on the
goal of being truly applied. The book is
geared toward speech-language patholo-
gists (SLPs) but should also be useful to
professionals in other disciplines, such as
ABA and special education.
Therefore, this textbook can be con-
sidered an interdisciplinary introduc-
tion to behavior analysis and contains
more than enough technical informa-
tion to challenge even the most serious
ABA student, including those pursuing
the credential of board certified behav-
ior analyst (BCBA). However, it is writ-
ten primarily to speak to those who are
in training for a career in communicative
disorders by including examples relevant
and pertinent to this discipline. In an ever-
changing and increasingly interdisciplin-
ary atmosphere, a textbook of this kind is
appropriate and timely.
The authors of this book represent
this professional diversity both in train-
ing and in specialized experience. The
authors include two doctoral-level SLPs,
a doctoral-level BCBA, and a combination
SLP/BCBA. All authors have experience
as university faculty (three in commu-
nicative disorders, one in psychology),
ix
and all authors have practical experience
working in clinical settings in a variety of
capacities. Prior to entering the university
setting, Christine A. Maul served as an
SLP in the public schools, with a specialty
in serving children and young adults with
severe developmental and intellectual dis-
abilities. She has been a faculty member in
the Department of Communicative Disor-
ders and Deaf Studies at California State
University, Fresno (CSUF) for 17 years,
as a lecturer and currently as an assistant
professor. As part of her duties, she super-
vises students performing clinical practi-
cum at the CSUF Speech, Language, and
Hearing Clinic, where individuals across
the life span with a wide range of commu-
nicative disorders are served. Brooke R.
Findley serves as an SLP in a rural school
district and part-time lecturer at CSUF.
After becoming an SLP, she recognized the
value in pursuing further education in the
field of ABA and is currently also certified
as a behavior analyst. Her areas of pro-
fessional interest include verbal behav-
ior, cultural and linguistic diversity, and
educational leadership. She is currently
pursuing a doctoral degree in Educational
Leadership at CSUF. Amanda Adams
started as a director for agencies provid-
ing autism services and then served as the
lead behavior analyst for a large school
district before accepting a faculty position
in the psychology department at CSUF. In
her 8 years at CSUF, she was the coordi-
nator for the ABA master’s program and
the founder of the Fresno State autism
center. She is now the CEO and executive
director of the California Autism Cen-
ter. In addition, another doctoral-level
SLP, Frances Pomaville, also an assistant
x  Behavioral Principles in Communicative Disorders: Applications to Assessment and Treatment
professor at CSUF, stepped forward to
write the chapter on verbal behavior to
help us meet our deadline. She did a thor-
ough job with a difficult topic, and we are
grateful for her contribution. We would
also like to thank graduate student assis-
tant Victoria Riley who gave us much help
in putting together the glossary.
The focus of this textbook is on
behavioral principles derived from learn-
ing theory, as discovered by the work
of behavioral scientists, chief among
them B. F. Skinner. The authors admit-
tedly share a significant bias toward the
behavioral viewpoint presented in this
text. Although we acknowledge the con-
tributions of other approaches, it was
our intent in writing this text to present
basic behavioral principles and propose
ways in which those principles can effec-
tuate positive change in communicative
behaviors. It is not within the scope of
the text to thoroughly examine the great
theoretically oriented debates that have
spun around behaviorism, particularly
the cognitive-linguistic versus the behav-
ioral viewpoints. Such theoretical discus-
sion regarding other approaches is largely
omitted from this text due to our desire
to maintain a clinically oriented focus on
behavioral principles.
We also acknowledge that bias against
behaviorism still exists in many modern-
day circles, although we find this to be
unfortunate. Criticisms regarding behav-
iorism were easier to understand before
the 1980s, before we had a body of research
supporting the applied extensions of the
basic and theoretical work resulting from
Skinner’s research. In the 1960s, the use
of behavioral techniques in applied set-
tings increased, improving outcomes in
people with mental and developmental
disabilities. Then, in the 1970s and 1980s,
the methodology to empirically validate
behavioral techniques progressed, and
the number of scientific publications dem-
onstrating the value of those techniques
consequently increased. Currently, the evi-
dence supporting the application of behav-
ioral principles to effectuate change in
human behavior is vast, and people seek-
ing to improve their communicative behav-
iors should have access to treatment that
incorporates those principles. In writing
this textbook, it was our intent to pre­sent
material specifically focused on empirically
validated behavioral techniques that have
shown efficacy in enhancing clinicians’
skills and clients’ outcomes.
In addition to our advocacy for the
science of behaviorism, we also share a
dedication to putting professional territo-
rialism behind us and embracing interdis-
ciplinary collaboration to provide the best
evidence-based services possible to the
people we serve. This is a highly personal
devotion among the authors of this book,
one of whom is the parent of a young man
with autism and mental illness. Participat-
ing in clinical intervention techniques not
only as a professional but also as a parent
may be one of the most profound ways
of experiencing the importance of sci-
entific collaboration in maximizing out-
comes for a loved one. This is a measure
of success that can be achieved only if
theoretical turf battles are abandoned in
favor of interdisciplinary collaboration
among professionals dedicated to pursu-
ing common goals within an evidence-
based framework. Our clinical fields are
experiencing a shift, and professionals
from complimentary disciplines (SLP,
school psychology, BCBA, special educa-
tion teachers, etc.) are working together
more than ever before. This is happen-
ing not only in public schools but also, to
an increasing degree, within the private
sector. Being knowledgeable about the
 xi
Preface  

valuable methods and techniques used
by our colleagues in neighboring fields of
practice is extremely helpful in strength-
ening our own practice as clinicians and,
more important, will be of benefit to the
clients we serve. We hope that you find
this book to be challenging, informative,
and useful in your professional growth
and clinical work.

—Amanda Nicolson Adams,

PhD, BCBA
California Autism Center,
Fresno, California
 Contributors

Amanda Nicolson Adams, PhD, BCBA

CEO, Executive Director
California Autism Center and Learning Group
Fresno, California

Brooke R. Findley, MA, CCC-SLP, BCBA

Lecturer
Department of Communicative Disorders and Deaf Studies
California State University, Fresno
Fresno, California

Christine A. Maul, PhD, CCC-SLP

Assistant Professor
Department of Communicative Disorders and Deaf Studies
California State University, Fresno
Fresno, California

Frances Pomaville, PhD, CCC-SLP

Assistant Professor and Speech-Language Pathologist
Department of Communicative Disorders and Deaf Studies
California State University, Fresno
Fresno, California
Chapter 2

xiii
To M. N. Hegde
 Chapter 1

Introduction to Behaviorism

Chapter Outline

n Why Should SLPs Learn the Principles of Behaviorism and

How to Apply Them?
n Definition and History of Behaviorism
n Early Behavioral Scientists
Pavlov (1849–1936)
n

n Watson (1878–1958)

n B. F. Skinner (1904–1990)

n Principles of Behaviorism
n Operant Conditioning and Respondent Conditioning
n Positive and Negative Reinforcement
n Differential Reinforcement
n Positive and Negative Punishment
n Stimulus Discrimination and Generalization
n Behavior Modification: The Early Days
n Philosophical Underpinnings
n Early Experimentation in Behavior Modification
n Applied Behavior Analysis
n Definition of ABA
n Who Are Board Certified Behavior Analysts (BCBAs)?
n How Can SLPs and BCBAs Collaborate?

1
2  Behavioral Principles in Communicative Disorders: Applications to Assessment and Treatment
Why Should SLPs Learn the
Principles of Behaviorism
and How to Apply Them?
Speech-language pathologists (SLPs) are
in the business of changing communi-
cative behaviors in the clients that they
serve. SLPs seek to establish new commu-
nicative behaviors in their clients, rehabil-
itate communicative behaviors that may
have been lost, strengthen and maintain
existing communicative behaviors, and,
often simultaneously, decrease commu-
nicative behaviors that are undesirable.
SLPs are ethically charged to engage in
evidence-based practice to achieve these
goals, taking into consideration scientific
evidence reported in peer-reviewed jour-
nals, the preferences of their clients, and
their own clinical expertise (American
Speech-Language-Hearing Association,
2005). The well-researched principles of
behaviorism, therefore, are highly relevant
to the field of speech-language pathology
because it is a science that has resulted
in an evidence-based, systematic set of
methods applied to modify the behaviors
of others. On that basis alone, it is benefi-
cial for SLPs to have knowledge of those
methods and understand how they can be
applied in the assessment and treatment
of their clients. However, there are other
good reasons why SLPs should know and
apply principles of behaviorism.
SLPs are ethically charged to engage
in _______-_______ ______________.
Clients served by SLPs often present
with accompanying behavioral difficul-
ties. In the adult population, behavior
problems are a frequently occurring effect
of stroke and traumatic brain injury (TBI).
Also, the comorbidity of speech-language
difficulties and social emotional behavior
disorders (SEBDs) in children has been
well documented. It has been reported
that 21.6% of the child and adolescent
populations in the United States exhibit
behaviors that warrant the diagnosis of
some type of psychiatric disorder (Carter
et al., 2010). Among children who are
diagnosed with language disorders, there
is a much higher prevalence of SEBD;
estimates by various researchers range
between 50% and 70% (Redmond & Rice,
1998). Therefore, SLPs working in all set-
tings are likely to encounter behavioral dif-
ficulties in a significant percentage of the
clients they attempt to help. Often behav-
ior problems must be addressed before
clients can fully benefit from treatment.
It is also becoming increasingly likely
for SLPs to be asked to take part in col-
laborative, multidisciplinary efforts to
help children overcome behavioral dif-
ficulties that may impede their access to
educational curriculum (Bopp, Brown, &
Mirenda, 2004). In particular, SLPs may
be called upon to work closely with board
certified behavior analysts (BCBAs) who
are experts in discovering and describ-
ing problem behaviors in children and
adults and then writing behavior plans to
address those problems. BCBAs have spe-
cial expertise in applied behavior analy-
sis (ABA), a systematic set of methods
designed to discover aspects of a person’s
environment that contribute to problem
behavior through functional behavior
assessment (FBA). In addition, BCBAs
are experts in applying ABA principles to
build and shape new behaviors that may
be nonexistent or deficient in clients who
do not necessarily exhibit problem behav-
iors. SLPs can therefore collaborate with
BCBAs in assisting clients with a wide
range of communicative difficulties, with
or without accompanying behavior disor-
ders. This chapter ends with a further dis-

cussion of the origin of applied behavior
analysis and professional collaboration
between SLPs and BCBAs.
Clients with behavioral difficulties
will benefit from collaboration
between SLPs and _______-_______
______________ ______________.
Finally, although principles of ABA
are more closely associated with assess-
ment and intervention for children with
autism and other severe disabilities, those
principles have much broader application
to the entire gamut of types of commu-
nicative disorders, levels of severity, and
behaviors that simply interfere with the
delivery of therapy. SLPs who understand
the principles of ABA can address prob-
lems as critical as helping a nonverbal
individual obtain a system of communi-
cation to as mild as helping a child stay in
seat during therapy.
In summary, SLPs should learn and
apply principles of behaviorism because
(a) there is a well-established evidence
base for the efficacy of methods based on
behaviorism, (b) many clients on SLPs’
caseloads are likely to exhibit challeng-
ing behaviors, (c) there is an increasing
need for SLPs to collaborate with BCBAs
in providing assessment and interven-
tion for children with behavior disorders,
and (d) the principles of behaviorism are
broadly applicable across types and levels
of severity of communicative disorders.
Definition and History
of Behaviorism
Behaviorism as a philosophy is grounded
in the principles of empiricism and deter-
minism. These principles are not compat-
Introduction to Behaviorism   3
ible with mentalism, another widely held
philosophy that emphasizes assumed in-
ternal processes such as thought and per-
ception as the key to understanding why
human beings behave the way they do.
Empiricism is the belief that knowl-
edge can be derived only from sensory
experiences — from that which can be
seen, heard, touched, tasted, or smelled.
Therefore, direct observation is inextri-
cably necessary in establishing a knowl-
edge base. It is a well-founded principle
of behaviorism that human behavior must
be defined in terms of that which can be
observed and measured. For example, it is
not acceptable to a behaviorist to describe
a behavior as an internal emotional event,
such as joy. A behaviorist would instead
describe and quantify behaviors that
might indicate the person is joyful, such
as smiling, laughing, jumping up and
down, or making cheering noises.
Empiricists also do not accept an idea
simply because it appears to be logical,
well argued, or coherent. They insist on
putting ideas to the empirical test, a scien-
tific experiment arranged to directly expe-
rience the truth or untruth of a statement
or hypothesis. Principles of behaviorism
have been thoroughly examined through
empirical research, most often utilizing
single-case research design methodol-
ogy, which is described in more depth in
Chapter 9.
The belief that knowledge can be
derived only from sensory experi-
ences is called ______________.
Determinism is the belief that nothing
that happens in the world is haphazard.
Determinists believe that there is a system-
atic order in which phenomena relate to
each other. Every effect has a cause, and it is
possible through scientific experimentation
4  Behavioral Principles in Communicative Disorders: Applications to Assessment and Treatment
to discover, understand, and exert control
over a cause-effect relationship.
The belief that every effect has a cause
is called ______________.
The philosophical concepts of empir-
icism and determinism demand explana-
tions of human behavior that cannot be
derived from a mentalistic viewpoint.
Mentalism is the belief that humans act
the way they do because of internal,
unobservable phenomena, such as mind,
thought, and free will. There are a myr-
iad of other hypothetical constructs that
mentalistic authors have set forth. Freud
(1949), for example, described id, ego,
and super ego as the prime determinants
of human behavior. Chomsky (1965)
proposed an internal language acquisi-
tion device (LAD) as the mechanism by
which children acquire language. Mod-
ern-day authors have proposed many
models showing how the brain processes
information in much the same way as a
computer does without the neurological
data to prove such processes (e.g., Schyns,
Gosslin, & Smith, 2009). All of these are
examples of hypothetical constructs that
do not lend themselves to observation
and measurement and therefore can-
not be subjected to empirical testing.
Behavioral scientists therefore argue that,
although such constructs may be a part of
the human experience, they do not con-
tribute much to a functional understand-
ing of why human beings behave the way
they do.
The belief that humans act the
way they do because of internal,
unobservable phenomena is called
______________.
Early Behavioral Scientists
Behavioral principles have been in opera-
tion since the beginning of the history of
humankind. In response to the conditions
of their environments, humans learn to
seek shelter when it is cold, to hunt and
to farm in order to obtain food, to run in
times of danger, to swim instead of drown,
and so forth. Parents teach their children
by praising them when they are behaving
in a desirable way and by punishing them
when they are not. People repeat behav-
iors that result in rewards such as a pay-
check from a boss, a kiss from a spouse, a
hug from a child, or a smile from a friend.
Although these are everyday occur-
rences that are taken for granted by the
general public, the way in which behav-
ioral principles shape our lives was not
fully described or understood until sci-
entists began to systematically study how
living organisms respond to environmen-
tal stimuli. Three scientists who contrib-
uted to the founding of behavioral sci-
ence were Ivan Pavlov (1849–1936), John
B. Watson (1878–1958), and B. F. Skinner
(1904–1990).
Pavlov (1849–1936)
To understand Pavlov’s pioneering work,
it is necessary to distinguish between an
unconditioned stimulus, which elicits
an unconditioned response, and a con-
ditioned stimulus, which elicits a condi-
tioned response. Unconditioned stimuli
automatically elicit an unconditioned
response; living organisms, including
human beings, react to certain stimuli in
certain ways from birth, and those reac-
tions persist throughout the life span. For
example, the presentation of good food
elicits salivation, touching a hot plate elic-
its a rapid removal of the hand, experienc-
 Introduction to Behaviorism   5

ing pain elicits moaning, pollen in the air
elicits sneezing, and so forth. In each of
these examples, there is an unconditioned
stimulus (such as pollen) and an uncondi-
tioned response (such as sneezing).
Pavlov established that it is pos-
sible to elicit an unconditioned response
to neutral stimuli that do not ordinarily
elicit such a response. In his classic experi-
ments, he presented a variety of neutral
stimuli, most famously the sound of a
bell, prior to the presentation of food to a
dog. The food, the unconditioned stimu-
lus, elicited the dog’s salivation. However,
after repeated pairings of the bell tone
with presentation of the food, the dog sali-
vated upon hearing the bell. At that point,
the tone became a conditioned stimulus
and the dog’s salivation became a condi-
tioned response (Figure 1–1).
Pairing a neutral stimulus to an uncon-
ditioned stimulus to eventually elicit an
unconditioned response by presentation
of the neutral stimulus alone is currently
referred to as respondent conditioning.
“Pavlov’s dog” became an icon for the
establishment of a set of behavioral tech-
niques based on respondent conditioning,
which, when applied ethically, can help
people overcome irrational fears, such as
fear of flying, or cease undesirable hab-
its, such as smoking or excessive alcohol
consumption. This type of respondent
conditioning, also called Pavlovian or
classical conditioning, has limited appli-
cation to the treatment of communica-
tive disorders. However, Display Box 1–1
shows an example of the importance of
understanding the concept of respondent
conditioning.
Watson (1878–1958)
Watson was the mentalist’s biggest foe. He
soundly denounced the idea that scientific
inquiry in the field of psychology should
be concerned with unobservable men-
tal states (e.g., thought, mood, dreams,
theoretical constructs of consciousness,
etc.). He proposed instead that studies be
conducted solely to discover the causes
of observable behavior, with the ultimate

FIRST THEN

Neutral Unconditioned Conditioned

Stimulus Paired with Stimulus Stimulus
(NS) (US) (CS; formerly NS)

By itself. . .
Elicits
Unconditioned
Response (UR) Elicits Conditioned
Response
(CR; formerly UR)

Figure 1–1. The process of respondent conditioning.

6  Behavioral Principles in Communicative Disorders: Applications to Assessment and Treatment

Display Box 1–1. Keep Those

White Coats in the Closet!

Although respondent conditioning has little application to the

treatment of communicative disorders, it is beneficial for cli-
nicians to understand how people can have reflexive, condi-
tioned reactions to stimuli that would not ordinarily elicit such
reactions. For example, fear responses from a child may be elic-
ited by a clinician wearing a white lab coat. Previous trips to
the doctor’s office, in which people often wearing white coats
administer vaccinations, may have conditioned the children
to react in such a manner. Similarly, the mere sight of a tongue
depressor may elicit a backward jerk of the head or even a gag
reflex in clients who have previously experienced oral facial
examinations.
Based on the schematic given in Figure 1–1, identify the
unconditioned stimuli and the unconditioned responses,
leading to the conditioned stimuli and conditioned responses
described here.

goal of devising methods by which
human behavior can be predicted and
controlled. His ideas formed the founda-
tion for behaviorism as a purely objective
natural science leading to understanding
the relationships between environmental
stimuli and responses of living organisms
to those stimuli.
Watson was a colorful character
with strong opinions that led him in vari-
ous directions, most notably into animal
behavior, childrearing, and marketing.
In his most controversial experiment, he
applied the Pavlovian principle of respon-
dent conditioning to an 11-month-old
child, dubbed “Little Albert,” presenting
a furry little white rat to the child simulta-
neously with the disturbingly loud noise
of an iron rod clanging (Watson & Rayner,
1920). Albert had exhibited no previous
fear of the rat, but he understandably
reacted with fear to the clang of the iron
rod. After several simultaneous presenta-
tions of the two stimuli, the child began
to exhibit fearful reactions to the presen-
tation of the rat alone and also to other
animals and objects that had any kind
of resemblance to the rat. Thus, Watson
proved that fear could be a conditioned
response that could generalize to related
objects. Watson did not attempt to de-con-
dition Little Albert, and because the little
boy died at the age of 6 from hydrocepha-
lus, it was not possible to determine what
effect this experiment might have had on
his life. This type of experimentation, cou-
pled with Watson’s penchant for strongly
worded hyperbolic statements, discred-
ited him and the newly proposed science
of behaviorism in the eyes of many. Dis-
play Box 1–2 presents one of Watson’s
most controversial statements.
 Introduction to Behaviorism   7

Display Box 1–2. Watson’s “Dozen Infants” Quote

Watson wrote in the area of childrearing, advocating for a type

of emotionally detached style of parenting in which children
would be regarded and treated as young adults. He was so
convinced of the possibilities of molding children through con-
trol of environmental stimuli that he went so far as to propose
a 20-year moratorium on pregnancies to give behavioral sci-
entists time to gather data to propose the most efficient way
to raise children. One of his most widely quoted statements
(Watson, 1928) indicated his fervent belief in the power of
behavioral parenting:

Give me a dozen healthy infants, well-formed, and my own

specified world to bring them up in, and I’ll guarantee to
take any one at random and train him to become any type
of specialist I might select — doctor, lawyer, artist, merchant-
chief and, yes, even beggar-man and thief. (p. 104)

For more information on Watson’s writings on childrearing,

refer to Houk (2000).

responses to environmental stimuli. He

In the Little Albert experiment, what
were the unconditioned stimulus,
the neutral stimulus, and the uncon-
ditioned response? What eventually
became the conditioned stimulus and
the conditioned response?
B. F. Skinner (1904–1990)
Behaviorism as a science did not exist until
B. F. Skinner addressed human behavior
as observable, measurable events that
could be explained through discovery of
the causes of those behaviors. Through
his experiments with laboratory animals,
mostly rats and pigeons, Skinner dis-
covered patterns of behavior caused by
devised the Skinner box, although Skin-
ner did not like the term Skinner box to
describe the device he invented. He pre-
ferred terms such as lever box or operant
conditioning chamber. Figure 1–2 provides
a schematic of a Skinner box.
The Skinner box was a cage contain-
ing a lever that, when pressed, would
dispense a morsel of food. A rat running
around the cage would eventually acci-
dentally depress the lever — a behavior
that was rewarded, or reinforced, by the
presentation of food. Skinner observed
that the rat’s lever-pressing behavior
would then increase, as a result of having
received the food. This learning process,
in which a behavior is increased through
a reinforcing event, is one component of
8  Behavioral Principles in Communicative Disorders: Applications to Assessment and Treatment
Figure 1–2. The Skinner box.
what Skinner called operant condition-
ing. This was but one of many principles
of behaviorism established through this
type of experimental manipulation of
antecedent and consequence events caus-
ing a change in animal behavior.
The learning process in which
a behavior is increased through a
reinforcing event is one component of
what Skinner called ______________
______________.
Principles of Behaviorism
Through many and repeated experiments,
Skinner demonstrated patterns of animal
behavior that are considered founda-
tional principles of behaviorism. These
broad principles are briefly described
here. In later chapters, these principles
are discussed in more detail and applica-
tions to clinical practice, with examples,
are given.
Operant Conditioning and
Respondent Conditioning
Operant conditioning is the primary
process through which Skinner believed
people learn to behave the way that
they do. Through operant conditioning,
behavior is shaped and maintained by
the consequences that immediately fol-
low the behavior. Consequences alter the
frequency of the occurrence of behavior
either by increasing the behavior through
reinforcement or decreasing, or often
entirely eliminating, a behavior through
punishment. From birth, human beings
encounter a myriad of consequences on
the way to becoming persons who behave
the way they do. A small infant may curl
his lips in a reflexive response to passing
Another random document with
no related content on Scribd:
 Fig. 136.—Portion of the
radula of Gadinia
peruviana Sowb., Chili. ×
250. Type (c).
(c) Radula with an indefinite number of marginals, laterals (if
present) merging into marginals, central tooth present or absent,
inconspicuous, teeth all very small. This type of radula, among the
Nudibranchiata, is characteristic of certain sub-genera of Doris (e.g.
Chromodoris, Aphelodoris, Casella, Centrodoris), of Hypobranchiaea
and Pleurophyllidia; among the Tectibranchiata, of Actaeon, many of
the Bullidae, Aplustrum, the Aplysiidae, Pleurobranchus, Umbrella
and Gadinia (Figs. 136 and 137, C).
In the Pteropoda there are two types of radula. The
Gymnosomata, which are in the main carnivorous, possess a radula
with a varying number (4–12) of sickle-shaped marginals, central
tooth present or absent. In the Thecosomata, which feed on a
vegetable diet, there are never more than three teeth, a central and
a marginal on each side; teeth more or less cusped on a square
base.
Pulmonata.—The radula of the Testacellidae, or carnivorous land
Mollusca, is large, and consists of strong sickle-shaped teeth with
very sharp points, arranged in rows with or without a central tooth, in
such a way that the largest teeth are often on the outside, and the
smallest on the inside of the row (as in Rhytida, Fig. 139). The
number and size of the teeth vary. In Testacella and Glandina, they
are numerous, consisting of from 30 to 70 in a row, with about 50
rows, the size throughout being fairly uniform. In Aerope they are
exceedingly large, and only eight in a row, the outermost marginal
being probably the largest single tooth in the whole of the Mollusca.
The central tooth is always obscure, being, when present, simply a
weaker form of the weakest lateral; in genera with only a few teeth in
a row it is generally absent altogether.

Fig. 137.—Portions of the radula of

Opisthobranchiata, illustrating
types (b) and (c); A, Scaphander
lignarius L.; A´, one of the teeth
seen from the other side, × 40;
B, Lamellidoris bilamellata L.,
Torbay, × 60; C, Hydatina physis
L., E. Indies, × 75.
The first family of jaw-bearing snails, the Selenitidae, is distinctly
intermediate. The possession of a jaw relates it to the main body of
Helicidae, but the jaw is not strong, while the teeth are still, with the
exception of the central, thoroughly Testacellidan. The central tooth
is quite rudimentary, but it is something more than a mere weak
reproduction of the marginals. There are no true laterals. The
Limacidae show a further stage in the transition. Here the central
tooth has a definite shape of its own, tricuspid on a broad base,
which is more or less repeated in the first laterals; these, as they
approach the marginals, gradually change in form, until the outer
marginals are again thoroughly Testacellidan.[326] This is the general
form of radula, varied more or less in different genera, which occurs
in Nanina, Helicarion, Limax, Parmacella, and all the sub-genera of
Zonites. It is certain that some, and probable that all of these genera
will, on occasion, eat flesh, although their usual food appears to be
vegetable. The jaw is more powerful than in the Selenitidae, but
never so large or so strongly ribbed as in Helix proper.

Fig. 138.—Portion of the radula of Glandina truncata

Gmel. × 40.
 Fig. 139.—Portion of the radula of Rhytida
Kraussii Pfr., S. Africa. × 25.
When we reach the Helicidae, we arrive at a type of radula in
which the aculeate form of tooth—so characteristic of the Agnatha—
disappears even in the marginals, and is replaced by teeth with a
more or less quadrate base; the laterals, which are always present,
are intermediate in form between the central and the marginals, and
insensibly pass into the latter. In size and number of cusps the first
few laterals resemble the central tooth; in the extreme marginals the
cusps often become irregular or evanescent. As a rule, the teeth are
set squarely in the rows, with the exception of the extreme
marginals, which tend to slope away on either side. In some
Helicidae there is a slight approximation to the Zonitidae in the
elongation of the first marginals.
The above is the type of radula occurring in the great family
Helicidae, which includes not only Helix proper, with several
thousand species, but also Arion, Bulimus, Ariolimax, and other
genera. The jaw is almost always strongly transversely ribbed.
In the Orthalicidae (Fig. 140, C) the teeth of the radula, instead of
being in straight rows, slope back at an angle of about 45 degrees
from the central tooth. The central and laterals are very similar, with
an obtuse cusp on rather a long stem; the marginals become
bicuspid.
In the Bulimulidae, which include the important genera
Placostylus, Amphidromus, Partula, Amphibulimus, and all the
groups of South American Bulimulus, the jaw is very characteristic,
being thin, arched, and denticulated at the edges, as if formed of
numerous narrow folds overlapping one another. The radula is like
that of the Helicidae, but the inner cusp of the laterals is usually
lengthened and incurved. In Partula the separation between laterals
and marginals is very strongly marked.
The remaining families of Pulmonata must be more briefly
described. In the Cylindrellidae there are three distinct types of
radula: (a) Central tooth a narrow plate, laterals all very curiously
incurved with a blunt cusp, no marginals (Fig. 140, D); (b) radula
long and narrow, central tooth as in (a), two laterals, and about eight
small marginals; (c) much more helicidan in type, central and laterals
obtusely unicuspid, marginals quite helicidan. Type (c) is restricted to
Central America, types (a) and (b) are West Indian.
Pupidae: Radula long and narrow; teeth of the helicidan type,
centrals and laterals tricuspid on a quadrate base, marginals very
small, cusps irregular and evanescent. This type includes Anostoma,
Odontostomus, Buliminus, Vertigo, Strophia, Holospira, Clausilia,
and Balea.
Stenogyridae, including Achatina, Stenogyra, and all its sub-
genera: Central tooth small and narrow, laterals much larger,
tricuspid, central cusp long, marginals similar, but smaller.
Achatinellidae: Two types occur; (a) teeth in very oblique rows,
central, laterals, and marginals all of the same type, base narrow,
head rather broad, with numerous small denticles (Achatinella
proper, with Auriculella and Tornatellina, Fig. 140, E); (b) central
tooth small and narrow, laterals bicuspid, marginals as in Helix
(Amastra and Carelia).
 Fig. 140.—Portions of the radula of A, Hyalinia nitidula Drap.,
Yorkshire, with central tooth, first lateral, and a marginal
very highly magnified; B, Helix pomatia L., Kent, showing
central tooth, laterals, and one extreme marginal, the two
former also highly magnified; C, Orthalicus undatus Brug.,
Trinidad, with three laterals highly magnified; D, Cylindrella
rosea Pfr., Jamaica, central tooth and laterals, the same
very highly magnified; E, Achatinella vulpina Fér., Oahu,
central tooth (c) and laterals, the same highly magnified.
Succineidae: Central and laterals helicidan, bi- or tricuspid on a
quadrate plate, marginals denticulate on a narrow base; jaw with an
accessory oblong plate.
Janellidae: Central tooth very small, laterals and marginals like
Achatinellidae (a).
Vaginulidae: Central, laterals, and marginals unicuspid
throughout, on same plan.
Onchidiidae: Rows oblique at the centre, straight near the edges;
central strong, tricuspid; laterals and marginals very long, falciform,
arched, unicuspid.
 Auriculidae: Teeth very small; central narrow, tricuspid on rather a
broad base; laterals and marginals obscurely tricuspid on a base like
Succinea.
Limnaeidae: Jaw composed of one upper and two lateral pieces;
central and lateral teeth resembling those of Helicidae; marginals
much pectinated and serriform (Fig. 141, A). In Ancylus proper the
teeth are of a very different type, base narrow, head rather blunt,
with no sharp cusps, teeth similar throughout, except that the
marginals become somewhat pectinated (Fig. 141, B); another type
more resembles Limnaea.
 Fig. 141.—Portions of the radula of
A, Limnaea stagnalis L., with
the central tooth and two first
laterals, and two of the
marginals, very highly,
magnified; B, Ancylus fluviatilis
Müll., with two of the marginals
very highly magnified; C, Physa
fontinalis L., with central tooth
and two of the marginals very
highly magnified.
Physidae: Jaw simple, but with a fibrous growth at its upper edge,
which may represent an accessory plate; radula with very oblique
rows, central tooth denticulate, laterals and marginals serriform,
comb-like, with a wing-like appendage at the superior outer edge
(Fig. 141, C).
 Chilinidae: Central tooth small, cusped on an excavated triangular
base, marginals five-cusped, with a projection as in Physa, laterals
comb-like, serrations not deep.
Amphibolidae: Central tooth five-cusped on a broad base, central
cusp very large; two laterals only, the first very small, thorn-like, the
second like the central tooth, but three-cusped; laterals simple,
sabre-shaped.
Scaphopoda.—In the single family (Dentaliidae) the radula is
large, and quite unlike that of any other group. The central tooth is a
simple broad plate; the single lateral is strong, arched, and slightly
cusped; the marginal a very large quadrangular plate, quite simple;
formula, 1.1.1.1.1 (Fig. 133, B).
Cephalopoda.—The radula of the Cephalopoda presents no
special feature of interest. Perhaps the most remarkable fact about it
is its singular uniformity of structure throughout a large number of
genera. It is always very small, as compared with the size of the
animal, most of the work being done by the powerful jaws, while the
digestive powers of the stomach are very considerable.
The general type of structure is a central tooth, a very few laterals,
and an occasional marginal or two; teeth of very uniform size and
shape throughout. In the Dibranchiata, marginals are entirely absent,
their place being always taken, in the Octopoda, by an accessory
plate of varying shape and size. This plate is generally absent in the
Decapoda. The central tooth is, in the Octopoda, very strong and
characteristic; in Eledone and Octopus it is five-cusped, central cusp
strong; in Argonauta unicuspid, in Tremoctopus tricuspid. The
laterals are always three in number, the innermost lateral having a
tendency to assume the form of the central. In Sepia the two inner
laterals are exact reproductions of the central tooth; in Eledone,
Sepiola, Loligo, and Sepia, the third lateral is falciform and much the
largest.
 Fig. 142.—Portion of the radula of Octopus
tetracirrhus D. Ch., Naples, × 20.
In Nautilus, the only living representative of the Tetrabranchiata,
there are two sickle-shaped marginals on each side, each of which
has a small accessory plate at the base. The two laterals and the
central tooth are small, very similar to one another, unicuspid on a
square base.

Fig. 143.—Alimentary canal of Helix aspersa L.: a, anus; b.d, b.d´, right and
left biliary ducts; b.m, buccal mass; c, crop; h.g, hermaphrodite gland; i,
intestine; i.o, opening of same from stomach (pyloric orifice); l, l´, right
and left lobes of liver; m, mouth; oe, oesophagus; r, rectum; s.d, salivary
duct; s.g, salivary gland; st, stomach; t, left tentacle. (After Howes and
Marshall, slightly modified.)
 Salivary glands are found in most Glossophora. They occur in one
or two pairs on each side of the pharynx and oesophagus, the duct
usually leading forwards and opening into the anterior part of the
pharynx (see Figs. 143, 144). They are exceptionally large in the
carnivorous Gasteropoda. In certain genera, e.g. Murex, Dolium,
Cassis, Pleurobranchus, the secretions of these glands are found to
contain a considerable proportion (sometimes as much as 4·25 per
cent) of free sulphuric acid. This fact was first noticed by Troschel,
who, while handling a Dolium galea at Messina, saw the creature
spit a jet of saliva upon a marble slab, which immediately produced a
brisk effervescence. A number of the genera thus provided bore
through the shells of other Mollusca and of Echinoderms, to prey
upon their soft tissues, and it is possible that the acid assists in the
piercing of the shell by converting the hard carbonate of lime into
sulphate of lime, which can easily be removed by the action of the
radula.[327] In the majority of the Cephalopoda there are two pairs of
salivary glands, one lying on each side of the mouth, the other on the
middle of the oesophagus.
Fig. 144.—Alimentary canal, etc.,
of Sepia officinalis L.: a,
anus; b.d, one of the biliary
ducts; b.m, buccal mass; c,
coecum; i, ink-sac; i.d, duct
of same; j, jaws; l.l, lobes of
the liver; oe, oesophagus; p,
pancreatic coeca; r, rectum;
s.g, salivary glands; st,
stomach. (From a specimen
in the British Museum.)
 Fig. 145.—Gizzard of
Scaphander lignarius L.: A,
showing position with regard
to oesophagus (oe) and
intestine (i), the latter being
full of comminuted fragments
of food; p, left plate; p´, right
plate; p.ac, accessory plate;
B, the plates as seen from
the front, with the enveloping
membranes removed,
lettering as in A. Natural
size.

Fig. 146.—Section of the stomach of Melongena,

showing the gastric plates (g.p, g.p,) for the
trituration of food; b.d, biliary duct; g.g, genital
gland; i, intestine; l, liver; oe, oesophagus; st,
stomach. (After Vanstone.)
 3. The Oesophagus.—That part of the alimentary canal which lies
between the pharynx and the stomach (in Pelecypoda between the
mouth and stomach) is known as the oesophagus. Its exact limits
are not easy to define, since in many cases the tube widens so
gradually, while the muscular structure of its walls changes so slowly
that it is difficult to say where oesophagus ends and stomach begins.
As a rule, the oesophagus is fairly simple in structure, and consists
of a straight and narrow tube. In the Pulmonata and
Opisthobranchiata it often widens out into a ‘crop,’ which appears to
serve the purpose of retaining a quantity of masticated food before it
passes on to the stomach. In Octopus and Patella the crop takes the
form of a lobular coecum. In the carnivorous Mollusca the
oesophagus becomes complicated by the existence of a varying
number of glands, by the action of which digestion appears to begin
in some cases before the food reaches the stomach proper.
4. The Stomach.—At the posterior end of the oesophagus lies the
muscular pouch known as the stomach, in which the digestion of the
food is principally performed. This organ may be, as in Limax, no
more than a dilatation of the alimentary canal, or it may, as is usually
the case, take the form of a well-marked bag or pocket. The two
orifices of the stomach are not always situated at opposite ends;
when the stomach itself is a simple enlargement of the wall of one
side of the alimentary canal, the cardiac or entering orifice often
becomes approximated to the orifice of exit (pyloric orifice).
The walls of the stomach itself are usually thickened and
strengthened by constrictor muscles. In some Nudibranchs
(Scyllaea, Bornella) they are lined on the inside with chitinous teeth.
In Cyclostoma, and some Bithynia, Strombus, and Trochus there is a
free chitinous stylet within the stomach.[328] In Melongena (Fig. 146)
the posterior end of the oesophagus is provided with a number of
hard plate-like ridges, while the stomach is lined with a double row of
cuticular knobs, which are movable on their bases of attachment,
and serve the purpose of triturating food.[329] Aplysia has several
hard plates, set with knobs and spines, and similar organs occur in
the Pteropoda. But the most formidable organ for the crushing of
food is possessed by the Bullidae, and particularly by Scaphander
(Fig. 145). Here there is a strong gizzard, consisting of several plates
connected by powerful cartilages, which crush the shells, which are
swallowed whole.
Into the stomach, or into the adjacent portions of the digestive
tract, open the ducts which connect with the so-called liver. The
functions of this important organ have not yet been thoroughly
worked out. The liver is a lobe-shaped gland of a brown-gray or light
red colour, which in the spirally-shelled families usually occupies the
greater part of the spire. In the Cephalopoda, the two ducts of the
liver are covered by appendages which are usually known as the
pancreatic coeca; the biliary duct, instead of leading directly into the
stomach, passes into a very large coecum (see Fig. 144) or
expansion of the same, which serves as a reservoir for the biliary
secretions. At the point of connexion between the coecum and
stomach is found a valve, which opens for the issue of the biliary
products into the stomach, but closes against the entry of food into
the coecum. In most Gasteropoda the liver consists of two distinct
lobes, between which are embedded the stomach and part of the
intestine. In many Nudibranchiata the liver becomes ‘diffused’ or
broken up into a number of small diverticula or glands connecting
with the stomach and intestine. The so-called cerata or dorsal lobes
in the Aeolididae are in effect an external liver, the removal of which
to the outside of the body gives the creature additional stomach-
room.
The Hyaline Stylet.—In the great majority of bivalves the
intestine is provided with a blind sac, or coecum of varying length.
Within this is usually lodged a long cylindrical body known as the
hyaline or crystalline stylet. In a well-developed Mytilus edulis it is
over an inch in length, and in Mya arenaria between two and three
inches. The bladder-like skin of the stylet, as well as its gelatinoid
substance, are perfectly transparent. In the Unionidae there is no
blind sac, and the stylet, when present, is in the intestine itself. It is
said to be present or absent indifferently in certain species.
The actual function performed by the hyaline stylet is at present a
matter of conjecture. Haseloff’s experiments on Mytilus edulis tend to
confirm the suggestion of Möbius, that the structure represents a
reserve of food material, not specially secreted, but a chemical
modification of surplus food. He found that under natural conditions it
was constantly present, but that specimens which were starved lost
it in a few days, the more complete the starvation the more thorough
being the loss; it reappeared when they were fed again. Schulze, on
the other hand, believes that it serves, in combination with mucus
secreted by the stomach, to protect the intestine against laceration
by sharp particles introduced with the food. W. Clark found that in
Pholas the stylet is connected with a light yellow corneous plate, and
imagined therefore that it acts as a sort of spring to work the plate in
order to comminute the food, the two together performing somewhat
the function of a gizzard.[330]
5. and 6. The Intestine, Rectum, and Anus.—The intestine, the
wider anal end of which is called the rectum, almost invariably
makes a bend forward on leaving the stomach. This is the case in
the Cephalopoda, Scaphopoda, and the great majority of
Gasteropoda. The exceptions are the bilaterally symmetrical
Amphineura, in which the anus is terminal, and many
Opisthobranchiata, in which it is sometimes lateral (Fig. 68, p. 159),
sometimes dorsal (Fig. 67). The intestine is usually short in
carnivorous genera, but long and more or less convoluted in those
which are phytophagous. In all cases where a branchial or
pulmonary cavity exists, the anus is situated within it, and thus varies
its position according to the position of the breathing organ. Thus in
Helix it is far forward on the right side, in Testacella, Vaginula, and
Onchidium almost terminal, in Patella at the back of the neck, slightly
to the right side (Fig. 64, p. 157).
In the rhipidoglossate section of the Diotocardia (Trochus, Haliotis,
etc.) the rectum passes through the ventricle of the heart, a fact
which, taken in conjunction with others, is evidence of their
relationship to the Pelecypoda.
 Fig. 147.—Ink-sac of
Sepia, showing its
relation to the rectum:
a, anus; d, duct of
sac; i.g, ink-gland; i.r,
portion of the sac
which serves as a
reservoir for the ink; o,
orifice of ink-gland; r,
rectum; sp, double set
of sphincter muscles
controlling upper end
of duct. (Modified from
Girod.)
In nearly all Pelecypoda the intestine is very long and convoluted,
being sometimes doubled forward over the mouth. Towards its
terminal part it traverses the ventricle of the heart, except in Ostrea,
Anomia, Teredo, and a few more. The anus is always at the posterior
end of the animal, adjacent to and slightly above the adductor
muscle.
Anal glands, which open into the rectum close to the anus, are
present in some Prosobranchiata, e.g. Murex, Purpura. In the
Cephalopoda the anal gland becomes of considerable size and
importance, and is generally known as the ink-sac (Fig. 147); it
occurs in all known living genera, except Nautilus. The ink-sac
consists of a large bag generally divided into two portions, in one of
which the colouring matter is secreted, while the other acts as a
reservoir for its storage. A long tube connects the bag with the end of
the rectum, the mouth of the tube being controlled, in Sepia, by a
double set of sphincter muscles.

The Kidneys

The kidneys, nephridia,[331] renal or excretory organs, consist

typically of two symmetrical glands, placed on the dorsal side of the
body in close connexion with the pericardium. Each kidney opens on
the one hand into the mantle cavity, close to the anus (see Fig. 64, p.
157), and on the other, into the pericardium. The venous blood
returning from the body passes through the vascular walls of the
kidneys, which are largely formed of cells containing uric acid. The
blood thus parts with its impurities before it reaches the breathing
organs.
The kidneys are paired in all cases where the branchiae are
paired, and where the heart has two auricles, i.e. in the Amphineura,
the Diotocardia (with the exception of the Neritidae), the Pelecypoda,
and all Cephalopoda except Nautilus, which has four branchiae, four
auricles, and four kidneys. In other Gasteropoda only one kidney
survives, corresponding to the left kidney of Zygobranchiate
Gasteropods.
Besides their use as excretory organs the kidneys, in certain
groups of the Mollusca, stand in very close relation to the genital
glands. In some of the Amphineura the generative products, instead
of possessing a separate external orifice of their own, pass from the
genital gland into the pericardium and so out through the kidneys
(see Fig. 61 C, D, p. 154). In the Diotocardia it is the right kidney
alone which serves, besides its excretory functions, as a duct for the
emission of the generative products, the left kidney being at the
same time greatly reduced in size. Thus in Patella the left
nephridium is small, the right being much larger; both function as
excretory organs, but the right serves as a mode of conveyance for
the seminal products as well. In certain Pelecypoda (e.g. Yoldia,
Avicula, Modiola, Pecten, Spondylus) the genital glands
communicate directly, and with a similar object, with the renal pouch
on the same side of the body, but in the majority of cases the orifices
are distinct.

The following memoirs will be found useful for further study of this
portion of the subject:—
D. Barfurth, Ueber den Bau und die Thätigkeit der
Gasteropodenleber: Arch. Mikr. Anat. xxii. (1883), pp. 473–524.
Th. Behme, Beiträge zur Anatomie und Entwickelungsgeschichte
des Harnapparates der Lungenschnecken: Arch. Naturges. iv. (1889),
pp. 1–28.
R. Bergh, Semper’s Reisen im Archipelago der Philippinen;
Nudibranchiata: Theil ii. Band ii. (1870–78), Band iii. (1880–1892).
W. G. Binney, Terrestrial Air-breathing Mollusks of the United
States: Bull. Mus. C. Z. Harv. iv. (1878), 450 pp.
„ On the Jaw and Lingual Membrane of North American
Terrestrial Pulmonata: Proc. Ac. Nat. Sc. Philad. (1875), pp. 140–243.
J. T. Cunningham, The renal organs (Nephridia) of Patella: Quart.
Journ. Micr. Sc. xxiii. (1883), pp. 369–375.
„ „ Note on the structure and relations of the kidney in
Aplysia: Mitth. Zool. Stat. Neap. iv. (1883), pp. 420–428.
R. von Erlanger, On the paired Nephridia of Prosobranchs, etc.:
Quart. Journ. Micr. Sc. xxxiii. (1892), pp. 587–623.
H. Fischer, Recherches sur la Morphologie du Foie des
Gastéropodes: Bull. Scient. France Belg. xxiv. (1892), pp. 260–346.
 C. Grobben, Morphologische Studien über den Harn- und
Geschlechtsapparat, sowie die Leibeshöhle, der Cephalopoden: Arb.
Zool. Inst. Wien, v. (1884), pp. 179–252.
„ Die Pericardialdrüse der Gasteropoden: ibid. ix. (1890), pp.
35–56.
B. Haller, Beiträge zur Kenntniss der Niere der Prosobranchier:
Morph. Jahrb. xi. (1885), pp. 1–53.
A. Hancock, On the structure and homologies of the renal organ in
the Nudibranchiate Mollusca: Trans. Linn. Soc. xxiv. (1864), pp. 511–
530.
A. Köhler, Microchemische Untersuchung der Schneckenzungen:
Zeits. Gesamm. Naturw. viii. (1856), pp. 106–112.
Ad. Oswald, Der Rüsselapparat der Prosobranchier: Jena. Zeits.
Naturw. N.F. xxi. (1893), pp. 114–162.
R. Perrier, Recherches sur l’anatomie et l’histologie du rein des
Gastéropodes prosobranches: Ann. Sc. Nat. Zool. (7), viii. (1889), pp.
61–315.
C. Semper, Reisen im Archipelago der Philippinen; Land
Pulmonata: Theil ii. Band iii. (1870–77).
C. Troschel, Das Gebiss der Schnecken: Berlin, 1856–1892.
W. G. Vigelius, Ueber das Excretionssystem der Cephalopoden:
Niederl. Arch. Zool. v. (1880), pp. 115–184.