Professional Documents
Culture Documents
Full Chapter Learning and Memory Basic Principles Processes and Procedures 6Th Edition W Scott Terry PDF
Full Chapter Learning and Memory Basic Principles Processes and Procedures 6Th Edition W Scott Terry PDF
https://textbookfull.com/product/boundary-retracement-processes-
and-procedures-1st-edition-donald-a-wilson/
https://textbookfull.com/product/principles-of-electron-optics-
basic-geometrical-optics-2nd-edition-peter-w-hawkes/
https://textbookfull.com/product/residential-construction-
academy-basic-principles-for-construction-5th-edition-mark-w-
huth/
https://textbookfull.com/product/cognition-6th-edition-scott-
sinnett/
Basic photographic materials and processes Fourth
Edition Josh Shagam
https://textbookfull.com/product/basic-photographic-materials-
and-processes-fourth-edition-josh-shagam/
https://textbookfull.com/product/cognitive-neuroscience-of-
memory-scott-d-slotnick/
https://textbookfull.com/product/associative-memory-cells-basic-
units-of-memory-trace-jin-hui-wang/
https://textbookfull.com/product/highland-lover-6th-edition-
amanda-scott/
https://textbookfull.com/product/research-methods-in-language-
acquisition-principles-procedures-and-practices-first-edition-
blume/
Learning and Memory
Learning and Memory provides a balanced review of the core
methods and the latest research on animal learning and human
memory. Topical coverage ranges from the basic and central
processes of learning, including classical and instrumental
conditioning and encoding and storage in long-term memory, to
topics not traditionally covered, such as spatial learning, motor skills,
and implicit memory. The general rules of learning are reviewed
along with the exceptions, limitations, and best applications of these
rules. Alternative approaches to learning and memory, including
cognitive, neuroscientific, functional, and behavioral, are also
discussed. Individual differences in age, gender, learning abilities,
and social and cultural background are explored throughout the text
and presented in a dedicated chapter.
The relevance of basic principles is highlighted throughout the text
with everyday examples that ignite reader interest in addition to
more traditional examples from human and animal laboratory
studies. Research examples are drawn from education,
neuropsychology, psychiatry, nursing, and ecological (or everyday)
memory. Each chapter begins with an outline and concludes with a
detailed summary. Applications and extensions are showcased in text
boxes as well as in distinct applications sections in every chapter;
and review and recapitulation sections are interspersed throughout
the chapters.
W. Scott Terry is Professor Emeritus at the University of North
Carolina at Charlotte where he taught psychology of learning to
undergraduates and graduate students in industrial-organizational
psychology, clinical psychology, and education. He received his Ph.D.
from Yale University, where he studied learning, cognitive
psychology, and psychobiology. He has published research in the
Journal of Experimental Psychology, Learning and Motivation,
Journal of Educational Psychology, and Teaching of Psychology. Over
the years, his research interests have included classical conditioning,
comparative cognition, personality and memory, and everyday lapses
of cognition.
Learning and Memory
Basic Principles, Processes, and
Procedures
Sixth Edition
W. Scott Terry
Designed cover image: © Getty Images
Sixth edition published 2023
by Routledge
605 Third Avenue, New York, NY 10158
and by Routledge
4 Park Square, Milton Park, Abingdon, Oxon, OX14 4RN
Routledge is an imprint of the Taylor & Francis Group, an informa business
© 2023 W. Scott Terry
The right of W. Scott Terry to be identified as authors of this work has been
asserted in accordance with sections 77 and 78 of the Copyright, Designs and
Patents Act 1988.
All rights reserved. No part of this book may be reprinted or reproduced or utilised
in any form or by any electronic, mechanical, or other means, now known or
hereafter invented, including photocopying and recording, or in any information
storage or retrieval system, without permission in writing from the publishers.
Trademark notice: Product or corporate names may be trademarks or registered
trademarks and are used only for identification and explanation without intent to
infringe.
First edition published by Pearson Education Inc. 2000
Fifth edition published by Routledge 2017
ISBN: 978-1-032-12973-0 (hbk)
ISBN: 978-1-003-22709-0 (ebk)
DOI: 10.4324/9781003227090
Typeset in Univers
by SPi Technologies India Pvt Ltd (Straive)
Access the Support Material: www.routledge.com/9781032129730
To:
Lorraine Piersanti Terry
Christopher Scott Terry
Jason Scott Terry
Brief Contents
Preface
Acknowledgments
1 Introduction
2 Habituation and Other Forms of Stimulus Learning
3 Classical Conditioning
4 Instrumental Conditioning: Reward
5 Instrumental Conditioning: Nonreward, Punishment,
and Avoidance
6 Verbal Learning
7 Human Memory Conceptual Approaches
8 Short-Term Memory
9 Encoding
10 Storage and Retrieval
11 Spatial, Motor-Skills, and Implicit Learning
12 Individual Differences in Learning and Memory
Glossary
References
Author Index
Subject Index
Contents
Preface
Acknowledgments
1 Introduction
The Origins of the Study of Learning
Epistemology
Evolution
Contemporary Influences
The Definition of Learning
Some Caveats
The Learning/Performance Distinction
Learning: Recapitulation
The Relationship Between the Terms Learning and Memory
The Science of Learning and Memory
Basic Research
Applied Research
Common Sense and Common Knowledge
Why Animals?
Conceptual Approaches to the Study of Learning
The Functional Approach
The Behavioral Approach
The Cognitive Approach
The Neuroscience Approach
Applications
Studying: Recommendations from Experts
Summary
2 Habituation and Other Forms of Stimulus Learning
What Is a Stimulus?
The Orienting Response
Habituation
Methods of Studying Habituation
Parametric Features of Habituation
Explanations of Habituation
Nonlearning Explanations
Neuroscience Theory
Evolution
Cognitive Theories
Sensitization
Perceptual Learning
Factors Affecting Perceptual Learning
Other Effects of Stimulus Exposure
Preference for Familiar Stimuli
Recapitulation: The Effects of Repeated Stimulus
Presentation
Applications
Exposure Therapy for Fear
Needle Fear
Summary
3 Classical Conditioning
The Definition of Classical Conditioning
Historical Note
Methods of Studying Classical Conditioning
Representative Procedures
What Stimuli Can Serve as CSs?
Basic Phenomena of Conditioning
Acquisition
Extinction
Generalization
Discrimination
Second-Order Conditioning
The Role of Contiguity
Summary of the Basic Phenomena
Other Factors Affecting Conditioning
Compound CSs
CS–US Relevance
Conditioned Inhibition
What Is Learned in Classical Conditioning?
So, What Is Learned in Pavlovian Conditioning?
The Role of Awareness in Conditioning
Extensions of Conditioning
Evaluative Conditioning
Conditioning with Drug USs and the Development of
Tolerance
Applications of Conditioning
The Conditioning Theory of Phobias
Overview of the Conditioning Theory of Phobias
Extinction as Therapy
Summary
4 Instrumental Conditioning: Reward
Definition and History
Thorndike and Trial-and-Error Learning
Skinner and Operant Learning
Elements of Instrumental Conditioning
Methods of Study
Positive Reinforcement
Reinforcement Variables Affecting Acquisition
Amount of Reinforcement
Drive
Schedules of Reinforcement
Delay of Reinforcement
Secondary Reinforcement
Social Reinforcement
Theories of Reinforcement
Drive Reduction
Incentive Motivation
The Neural Basis of Reinforcement
Reinforcers as Behaviors
Reinforcers as Strengtheners
Reinforcers as Information
So, What Is Reinforcement?
Is Reinforcement Necessary for Learning?
Awareness in Human Instrumental Learning
Criticisms of the Use of Reinforcement
Response Learning
Shaping
Limitations of Response Learning
Discriminative Stimulus Control
Generalization and Discrimination
Summary of Response Learning and Stimulus Learning
What Is Learned in Instrumental Conditioning?
Response–Reinforcer Learning
Stimulus–Response Learning
Stimulus–Reinforcer Learning
What Is Learned? Stimulus–Response–Reinforcer
Applications
Habits
Behavior Modification
Summary
5 Instrumental Conditioning: Nonreward, Punishment, and
Avoidance
Defining the Contingencies: Nonreward, Punishment, and
Avoidance
Extinction
The Partial Reinforcement Extinction Effect
Punishment
When Does Punishment Work?
Side Effects of Punishment
Punishment or Nonreward?
Persistence
Avoidance Learning
Theories of Avoidance Learning
Approach–Avoidance Conflict
Approach or Avoidance as a Coping Response
Prediction, Control, and Helplessness
Extensions of the Learned-Helplessness Concept
Summary of Learned Helplessness
Neuroscience and Aversive Learning
The Amygdala and Aversive Learning
Avoidance Conditioning
Applications of Aversive-Learning Contingencies
Pet Containment Systems
Treatment of Obsessive-Compulsive Disorder
Summary
6 Verbal Learning
The Ebbinghaus Legacy
Serial Learning
Serial Position
The Ubiquitous Serial-Position Curve
Remote Associations
Serial Learning: A Summary
Paired-Associate Learning
Analysis of Paired-Associate Learning
Direction of Associations
Paired-Associate Learning: A Summary
Free Recall
Serial-Position Effects
Rehearsal
Organization
Free Recall: A Summary
Recognition Memory
Recognition: Remembering Versus Knowing
Accessible Memories
Recall Versus Recognition Versus Relearning
Implicit Learning
Relationships Among the Verbal-Learning Tasks
Statistical Learning
Propositional Learning
Statistical Language Learning
Applications
Assessment of Cognitive Impairment
Mnemonics
Summary
Recognition Memory
7 Human Memory Conceptual Approaches
Partitioning Memory
Components of Memory Approach
Dual-Store Theory: Short-Term and Long-Term
Memories
Divisions of Long-Term Memory: Episodic and Semantic
Divisions of Long-Term Memory: Procedural Learning
and Priming
The Organization of Long-Term Memory
Stages of Memory Approach
Dissociating Stages
Processing Approaches to Memory
Depth of Processing
Artificial Neural Networks
Modeling Configural Learning
Connectionism and the Other Approaches
Applications
The Study of Abnormal Memory: Amnesia
Types of Amnesias
Everyday Forgetting and the Models of Memory
Summary
8 Short-Term Memory
Some History
Short-Term Memory Tasks
The Brown–Peterson Distractor Task
Memory Span
Characteristics of Verbal Short-Term Memory
Acoustic Encoding
Limited Capacity
Limited Duration
Forgetting: Short-Term Memory Is Sensitive to
Disruption
Transfer to Long-Term Memory
Control Processes
Summary of the Features of Short-Term Memory
Other Modalities of Short-Term Memory
Visual Short-Term Memory
Spatial Short-Term Memory
Short-Term Memory for Actions
Short-Term Memory for Odors
Short-Term Memory in the Hearing Impaired
Working Memory
Theoretical Overview of Working Memory
Measuring Working Memory
Executive Functions
Working Memory and Consciousness?
Working Memory and Culture
Individual Differences in Working Memory
Aging and Working Memory
Dementia and Working Memory
Anxiety and Working Memory
Multitasking
Is There Really a Separate Short-Term Memory?
A Single-Memory Approach?
Neuropsychological Dissociations of Two-Memory
Systems
One Memory or Several?
Applications
Comprehending and Using Language
Problem Solving
Summary
9 Encoding
Separating Encoding from Retrieval
Some Basic Variables in Encoding
Elaborative Rehearsal
What Exactly Is Elaborative Processing?
Imagery and Memory for Pictures
Meaningfulness
Presentation Variables
Testing Effects
Isolation Effects
Spacing Effects
Learner Variables
Incidental Versus Intentional Learning
Incentives
Arousal
Emotions and Encoding
Schemas
Metamemory
Encoding: Summing Up
Applications
Academic Learning and Encoding
Summary
10 Storage and Retrieval
Storage
Long-Term Memory for Naturalistically Learned Material
The Nature of Storage
Retrieval
Retrieval from Episodic Memory
What Makes a Good Retrieval Cue?
Emotional Arousal and Retrieval
Prospective Memory
Metamemory and Partial Retrieval
Feeling of Knowing
False Retrieval
False Memory
Source Memory
The Effect of Postevent Information
Debunking Misinformation
Recovered Memory
Retrieval Versus Reconstruction
Applications
Strategies for Searching Memory
Context-Specific Learning
Reconsolidation
Summary
11 Spatial, Motor-Skills, and Implicit Learning
Spatial Learning
Rats, Mazes, and Psychology
Routes Versus Cognitive Maps
Maze Learning and the Brain
Schemas in Spatial Memory
The Development of Spatial Memory in Children
Motor-Skills Learning
Practice
Knowledge of Results
Long-Term Retention of Skills
Implicit Learning
Some Implicit-Learning Tasks
Dissociating Categories of Implicit Learning
Expertise
From Declarative to Procedural
Applications
Implicit Learning
Spatial Memory
Developing Memory Skill
Summary
12 Individual Differences in Learning and Memory
The Nature of Nurture: The Genetics of Learning Ability
Animal Studies
Human Studies
Age Differences in Learning and Memory
Conditioning
Memory Development in Children
Aging and Memory
Exceptional Memory: The Mnemonists
Highly Superior Autobiographical Memory
Gender and Cognitive Abilities
Learning Styles
The Visualizer–Verbalizer Dimension
Kolb
Sternberg
Learning Styles?
Self-Control
Social and Cultural Differences
Epic Memories
Experimental Studies
Contemporary Cultural Psychology
Summary
Glossary
References
Author Index
Subject Index
Preface
Learning and remembering what we have learned are fundamental
psychological processes. The scientific study of learning and memory
has produced a sizable body of principles, laws, and heuristic rules.
Since the very beginning of psychology as a discipline, researchers
who study learning have been asked to provide instruction in these
principles. What do we know about learning, and how can this
knowledge be usefully applied? My approach to writing a book on
learning and memory is to present the basic methods and results of
our research and to emphasize the relevance of our research to
other disciplines.
My purpose in writing this book has remained the same since the
first edition. I wanted to write what I teach in my own courses. I
have taught Psychology of Learning to college undergrads, Master’s
level I/O psychology, and students in Education Master’s program. I
had a unique background that coincided with the resurgence in
behavioral Learning Theory, the development of Cognitive
Psychology, and new discoveries in the Biopsychology of Learning.
I had several goals in mind in writing this book. The topical
coverage is restricted to basic and central processes, such as
classical and instrumental conditioning, and encoding and storage in
long-term memory. Yet today, the basic processes should include
implicit memory, spatial learning, and remembering in the world
outside the laboratory. In addition to presenting some general rules
for learning, it is just as important to specify the exceptions and
limitations to those rules. When is spacing practice or giving
immediate reinforcement better, and when might massed practice or
delayed reward be better?
I have tried to write a book that will appeal to a broad audience
by including a range of research examples, from education,
neuropsychology, psychiatry, nursing, marketing, and ecological (or
everyday) memory. I value basic research on learning and memory.
This includes laboratory studies, often of animals, as fundamental
means for determining the principles by which learning occurs and
memory persists. Yet in my own teaching, I have found that the
relevance of these basic principles needs to be made clear
consistently. Lecturers often use everyday examples to maintain
student interest. I believe that a more important purpose is to
illustrate how basic principles can be translated into applications. To
be skilled and knowledgeable in the basics is not enough. For
instance, experts in animal conditioning are not necessarily good dog
trainers. (Just ask my dogs). We also need to consider the extent to
which our laboratory results will generalize to other subject
populations that differ in age, learning abilities, cultural background,
and even species (!).
A book on learning should also include aids to facilitate learning.
Each chapter begins with an outline and concludes with a detailed
summary. Review and recapitulation paragraphs are interspersed in
the chapters. Some topics and terms are repeated across chapters.
Essential terms are in boldface, and these also appear in the
glossary at the end of the book. Other important terms are in italics.
An instructor’s website will be available, to include lecture outlines,
additional material for use in class, suggested readings, student
activities, and links to online videos. A separate set of test questions
is also available.
Acknowledgments
A number of people have influenced me, either directly or indirectly,
in writing this book. I had the good fortune to have had mentors
both in college and graduate school. At Fairfield University, the
psychology faculty were enthusiastic, devoted, and student oriented.
They determined my choice of a career as a teacher and researcher
in the field of learning. At Yale University, the freedom to pursue
individual interests, and close faculty interactions, offered an exciting
environment in which to learn. Allan Wagner, in his classroom and in
his laboratory, became an enduring role model and mentor. (A note
to students: My guess is that many of your professors and
instructors had similar relationships that led to their own decisions to
become teachers. Ask them about it).
I would also like to thank my extended Terry family for
encouragement, patience in living with an absent-minded professor,
and leaving me alone for long periods of time. The next generation
of Terry children have renewed my wonder in the potential for
learning. A wise professor once said that if you want to learn about
Learning, have children. I’m still learning.
I would like to acknowledge the editorial and production staff at
Taylor & Francis / Routledge for their dedicated and professional
work on this book. Special thanks to Georgette Enriquez, psychology
editor at Taylor & Francis, who made this new edition possible. I
appreciate her encouragement and support. Thanks also to senior
editorial assistant Lakshay Gaba. The production staff corrected my
errors and updated the text’s design. Thank you senior production
editor Kris Šiošytė, and copyeditor Lorraine Savage, proofreader
Sandra Stafford, and indexer Merideth Murray.
Several university and college instructors commented on chapters
and successive drafts of the several editions of this book, providing
extensive commentary and their own insights into this field. For their
efforts, I am both grateful and respectful. Any errors and mistakes
that remain are mine, and the reviewers can safely reply, “I told you
so.”
Finally, I would like to thank my students, both undergraduate and
graduate, from the last too-many years, for letting me try out ideas
on them. I have found that if you present things in a good-natured
manner, students will respond similarly. They have been a valuable
source of feedback.
Chapter 1
Introduction
DOI: 10.4324/9781003227090-1
Contents
The Origins of the Study of Learning
Epistemology
Evolution
Contemporary Influences
The Definition of Learning
Some Caveats
The Learning/Performance Distinction
Learning: Recapitulation
The Relationship Between the Terms Learning and
Memory
The Science of Learning and Memory
Basic Research
Applied Research
Common Sense and Common Knowledge
Why Animals?
Conceptual Approaches to the Study of Learning
The Functional Approach
The Behavioral Approach
The Cognitive Approach
The Neuroscience Approach
Applications
Studying: Recommendations from Experts
Summary
Students who take a course in the psychology of learning are usually
knowledgeable about learning by the time they have reached this
point. Through instruction and on-the-job training, they have already
picked up numerous everyday, common sense principles about how
to learn. Students can readily tell their instructors that it is better to
spread studying over several days rather than to cram it all into one
day (what psychologists call the spaced versus massed practice
effect), and that temporary forgetting for otherwise well-known
information occurs, especially on exam days (what we otherwise call
retrieval failure). Students are aware of what psychologists call
context-dependent learning, that it is better to study in the place in
which you will take the test. These practical principles are accurate
as broad generalities, but they are also only partially true. They are
half-truths. This is a book full of half-truths.
Let me quickly explain what I mean. There are numerous facts,
laws, and principles of learning that have been uncovered over the
past 120 years that psychology has formally been in existence.
However, these principles are more complex than the simple
statements we popularly use to describe them (e.g., spaced practice
is better than massed practice). Statements of these principles
almost always require qualifiers; they are true under certain
conditions. In this book, I will attempt to tell both halves, and thus
in the end something closer to the truth as we know it now.
Take some well-known popular generalizations. Spaced practice
produces better learning than does massed practice. Well, yes,
usually. But much depends on what we are attempting to learn and
how long we will have to retain it, just two of several possible
qualifiers we might add here. Actually, one line of research on
remembering people’s names suggests that it is better to mass
repetitions of a given name at first, and then gradually lengthen the
interval between successive presentations (Landauer & Bjork, 1978).
(Note to readers: The standard format for noting sources in
psychology is to list the authors’ last name and the year of
publication. Complete source information is provided in the
References at the end of the book). Repeat the new name
immediately; repeat it again after a little while; and keep increasing
the interval to the next repetition. As a second example, common
sense seems to say that feedback is more effective when it is given
immediately and consistently after each performance of a behavior.
Yet, this is not always so. Skilled movements are sometimes learned
better with delayed or only occasional feedback (see Chapter 11).
Other forms of learning pose questions that have alternative
correct answers. Do subliminal audio messages, such as suggestions
to induce self-control or weight loss, work? Both yes and no answers
can be defended. Some data indicate they are effective, but more
probably due to a placebo effect. Does this mean that there is no
such thing as subliminal learning? No, learning can occur at many
levels of awareness or consciousness. Does sleep learning occur?
Instead of buying the hard-copy version of this text, should you get
the audio version to listen to throughout the night? The answer
depends on what you mean by learning. Research conducted in
sleep labs indicates that factual information is probably not being
learned, but possibly some other forms of learning (such as
conditioning of the Pavlov variety) might occur.
The point of these examples is to give a sample of what real
principles of learning look like. The goal of this book is to present a
scientifically accurate and sophisticated view of the principles of
learning. This includes the qualifying statements: when a given
principle holds and when exceptions occur. We can simplify a
description to aid comprehension, but it can be simplified only so
much before it becomes inaccurate.
Epistemology
The nature–nurture question asks how we are affected by biology on
the one hand (i.e., nature) and by environment on the other (i.e.,
nurture). If a child were raised in isolation from others, what would
that child know? Would the child instinctively develop into a kind and
just person, or one who is cruel and selfish? The the French
philosopher and social commentator Jean Rosseau (1712–1788) was
wary of the degrading influences of civilization, and believed that
“noble” peoples would be discovered living beyond the reach of so-
called civilization. However, the discovery of feral children, living
apart from other humans, were intellectually and emotionally
disabled (Candland, 1993).
The branch of philosophy known as epistemology studies how
we come to have knowledge. This is also the central question for the
field of learning. One view is that some knowledge is innate or
instinctive. Humans were controlled by instinctive motives such as
sociability, aggression, competitiveness, or protectiveness of family.
Further knowledge was developed by reasoning as illustrated by the
derivation of geometric axioms and algebraic logic. In each case,
knowledge is present independent of particular experiences with the
world.
By contrast, the British philosopher John Locke (1632–1704) said
that the origin of knowledge is in experience, as provided to the
mind through the senses. This is the philosophy of empiricism. For
instance, our notion of cause and effect is learned by our frequent
experiences in which one event in the world is regularly followed by
another event. Such regularities lead to the conclusion that the first
event is the cause of the second.
Empiricism led psychologists to investigate how we acquire
knowledge through experience. What is the origin of our association
from STOP to GO or TABLE to CHAIR? Locke said tables and chairs
are contiguous: They occur together in time or space. Beyond that,
tables and chairs are frequently contiguous. Therefore, our mental
representations of these separate ideas become associated in our
minds.
Early 20th century psychologists were receptive to the study of
association learning. For example, by adopting Ivan Pavlov’s method
of conditioning, experimenters paired a tone with food several times,
Another random document with
no related content on Scribd:
stylet will form the anterior tip of the proboscis, and will there be in a position for
offence or defence (Fig. 57, s).
In two out of the three groups into which the Nemertines are divided, the lateral
nerve-cords are in connexion with a network or plexus of nerves lying between
the muscular layers of the body-wall (Fig. 52, n.l), and in some forms constituting
a comparatively thick layer. In these two groups there are no definite nerve
branches except the anterior ones to the head. In the third group of Nemertines
the lateral nerve-cords lie within the muscular layers of the body-wall, and in this
case paired nerve branches are given off at definite intervals throughout the
whole length of the body. These branches divide up among the organs to which
they pass, and no nerve plexus is present.
The lateral cords vary in position in different cases. Sometimes they lie laterally,
at others the cords tend to approximate to one another in the median dorsal or in
the median ventral line, though in every case they remain distinctly separated.
Sense Organs.—Sense organs are usually present in the form of eyes arranged
at the sides of the head (Fig. 51, e), sometimes as a single pair and sometimes
in one or more groups on each side. The structure of the eyes varies from a
simple pigment spot to an organ which receives a special nerve-supply from the
brain, and possesses a refracting body answering to a lens, and behind this a
pigment layer and a layer of rods. Some forms are devoid of all traces of eyes.
Fig. 55.—Diagram to show the relations of the nervous system, circulatory system,
and proboscis sheath in the anterior end of the body in the Hoplonemertea,
modified from M‘Intosh. a.n, Nerves to anterior part of body and eyes; d.c,
dorsal commissure; d.n, median dorsal nerve; d.v, dorsal vascular trunk; l.v,
lateral vascular trunk; n.c, lateral nerve-cord; n.g.d, dorsal lobe of nerve
ganglion; n.g.v, ventral lobe of nerve ganglion; p.p, proboscis pore; p.s,
proboscis sheath; v.c, ventral commissure; v.s, vascular ring or collar.
Frontal Organ.—In many Nemertines there is present at the anterior tip of the
head a disc-shaped group of cells bearing long hairs or bristles. On this disc
open the secreting ducts of a number of gland cells lying in the head. It seems
possible that this frontal organ may function as an organ of taste.
In many forms the nervous system is charged with haemoglobin, which gives to
it a bright red colour.
The blood is usually colourless, but in some cases the corpuscles are coloured
red by haemoglobin.
Generative System.—The Nemertines are for the most part dioecious, only a
few certainly hermaphrodite species having been described, e.g. Tetrastemma
("Borlasia") kefersteinii Mar.[140]
The generative products in both cases are contained in sacs (Figs. 52, 53, g)
which lie in the lateral region of the body between the pouches of the alimentary
canal. The ova and spermatozoa are conveyed to the exterior by short ducts.
Most species are oviparous, though a few viviparous species are known (e.g.
Prosorhochmus claparedii).
For this reason Hubrecht divided the Nemertinea into three Orders—
Hoplonemertea, Schizonemertea, Palaeonemertea; the first of these Orders
corresponding with the Enopla, and the other two with the Anopla.
Order I. Hoplonemertea.
Fig. 58.—Head end of Cerebratulus marginatus Ren., from the ventral surface.
Drawn from a spirit specimen. Naples. × 1. c.s, Cephalic slit; m, mouth; p.p,
proboscis pore.
Principal British genera and species:—
The proboscis is unarmed. The epidermis and connective tissue form one layer,
below which is the basement membrane. The muscular layers are three in
number, two circular separated by a longitudinal layer. The nerve-cords lie
altogether external to the muscular layers, and are connected together
throughout by a plexus. No nerve branches are given off. The brain is not
divided into lobes. The cephalic slits are only represented by a shallow
depression on each side of the head, and no canals have been observed
leading from them. The intestine is straight, and the pouches are usually absent
or rudimentary. The circulatory system is largely made up of lacunar spaces, the
closed system being but little developed.
The Pilidium (Fig. 60) is a helmet-shaped larva bearing a tuft or spike dorsally,
and prolonged downwards laterally into a pair of lobes. The whole larva is
covered with cilia, there being a specially strong band round its ventral surface.
The dorsal spike is composed of a bunch of strongly developed cilia or of a long
flagellum. The alimentary canal consists of a sac constricted into oesophageal
and gastric regions (Fig. 60, oes and st). In this condition the larva swims about
freely in the water. The helmet-shaped Pilidium-skin forms no part of the future
Nemertine, the skin of which is developed as ingrowths from it; these meet one
another and unite to form a complete covering round the alimentary canal; the
larval skin is then cast off, and by a series of gradual steps the embryo develops
into the adult.
Habits.—Nemertines are often found under stones between high- and low-water
marks, lying on sandy or muddy bottoms. They are usually in the form of coiled
masses, and are generally in a state of quiescence. Hence it is probable that
their period of activity is during high-water, and that when left by the receding
tide they subside into a resting condition.
The large kinds, such as Lineus marinus, seem to be always found living alone,
but some of the smaller kinds, notably Tetrastemma dorsale and Prosorhochmus
claparedii, have gregarious habits and live in masses, the coils of the different
individuals being inextricably mixed.
Nemertines are commonly dredged from a depth of six or eight fathoms. They
may sometimes be found floating on the surface of the water, and some possess
the power of swimming rapidly, propelling themselves by a lateral motion of the
tail, the sides of which are in such cases prolonged into a thin fin-like edge. This
mode of progression is usually adopted by those which frequent deep water. A
pelagic Nemertine (Pelagonemertes) was discovered by Moseley near the
southern verge of the South Australian current, being found in a trawl with deep-
sea forms from a depth of 1800 fathoms. This animal was leaf-like in shape,
bluntly pointed behind and rather square in front.
Most Nemertines can be very readily kept in confinement. The chief apparent
effect of such a life is a loss of colour, the animal gradually becoming pallid in
hue. Owing also to the absence of proper food they diminish very much in size,
though even when all food is kept away an animal will sometimes continue to
live as long as eighteen months.
Owing to the force with which it is shot out, the proboscis is often completely
severed from the body, and in such a case the animal grows a new one in an
extremely short space of time. The proboscis thus broken off retains its power of
movement and contractility for a considerable time, and has been more than
once mistaken for a worm. This great vital power is probably due to the great
development of nervous tissue, the proboscis being usually richly supplied with
nerve plexuses.
In the armed Nemertines the eggs are deposited separately, and are not
connected together except by such accidental mucus as the animal deposits
normally; but in the unarmed a special mucous secretion forms a thick
investment for the eggs.
M‘Intosh[145] has observed the process of the deposition of the male and female
products in Nemertes gracilis. He put into a glass vessel a male and female of
this species in which the products were apparently ripe. Soon spermatozoa
began to issue in wreath-like jets from the body of the male, at first from the
middle region of the body, and afterwards anteriorly and posteriorly, until the
animal was enveloped in a dense cloud of spermatozoa. The whole process only
lasted a few minutes. When all the spermatozoa had apparently been given out,
the female was seen to protrude her head from the sand; she then passed to the
side of the vessel and deposited a group of eggs about three inches distant from
the spermatozoa.
The species found by Semper, and called by him Geonemertes palaensis, lives
under damp leaves and the roots of trees on Pelew Island in the North Pacific. It
is about 2 inches long, of a reddish-white colour, with narrow, brownish-black,
longitudinal stripes on its dorsal surface. It possesses six eyes and very small
cephalic slits and cerebral organs. The proboscis is armed, and opens by the
mouth instead of by a special pore.
They are probably more numerous than is at present known, and are
certainly scattered widely over the face of the earth, since they have
been found in Nicaragua, at Tashkend in Turkestan, and at
Philadelphia and Monroe in the United States.
The features which are supposed to indicate this are the elongated
vermiform shape showing no external signs of segmentation; the
ciliated smooth skin and the possession of unicellular mucous
glands; and the protrusible proboscis, which may be comparable to
the non-retractile proboscis of Balanoglossus, a comparison which is
strengthened by the fact that in some Nemertines a sheath of nerve-
fibres exists in the wall of the proboscis corresponding to the nerve
plexus in the proboscis of Balanoglossus. In both cases an
ectodermic nerve plexus exists with local thickenings along definite
lines, although these lines are not the same in the two cases. Both
possess a straight alimentary canal, ending in a terminal anus and
thrown out into paired lateral caeca, between which are the paired
metamerically-arranged generative sacs.
NEMATHELMINTHES & CHAETOGNATHA
BY
CHAPTER VI
NEMATHELMINTHES
INTRODUCTION—NEMATODA—ANATOMY—EMBRYOLOGY—
CLASSIFICATION—ASCARIDAE—STRONGYLIDAE—TRICHOTRACHELIDAE—
FILARIIDAE—MERMITHIDAE—ANGUILLULIDAE—ENOPLIDAE—PARASITISM
—NEMATOMORPHA—ANATOMY—CLASSIFICATION—LIFE-HISTORY—
ACANTHOCEPHALA—ANATOMY—EMBRYOLOGY—CLASSIFICATION.
i. The Nematoda.
ii. The Nematomorpha (Gordiidae).
iii. The Acanthocephala.
With few exceptions the shape of the body is filiform (Figs. 66 and
71), the two ends being more or less pointed, and the posterior end
of the male, which is generally a smaller animal than the female, is
usually slightly recurved. The worms are, as a rule, white, or of the
colour of polished ivory; they may be opaque or semi-transparent,
but pigment spots are rarely developed.