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 Third Edition

INTRODUCTORY
IMMUNOLOGY

Basic Concepts for

Interdisciplinary Applications

JefFrey K. Actor
Introductory Immunology
Basic Concepts for Interdisciplinary Applications
Introductory
Immunology
Basic Concepts for Interdisciplinary
Applications

Third Edition

Jeffrey K. Actor
Department of Pathology and Laboratory Medicine,
University of Texas McGovern Medical School, Houston, TX, United States
Academic Press is an imprint of Elsevier
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Copyright © 2023 Elsevier Inc. All rights reserved.
No part of this publication may be reproduced or transmitted in any form or by any means,
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arrangements with organizations such as the Copyright Clearance Center and the Copyright
Licensing Agency, can be found at our website: www.elsevier.com/permissions.
This book and the individual contributions contained in it are protected under copyright by the
Publisher (other than as may be noted herein).

Notices
Knowledge and best practice in this field are constantly changing. As new research and
experience broaden our understanding, changes in research methods, professional practices, or
medical treatment may become necessary.
Practitioners and researchers must always rely on their own experience and knowledge in
evaluating and using any information, methods, compounds, or experiments described herein.
In using such information or methods they should be mindful of their own safety and the safety
of others, including parties for whom they have a professional responsibility.
To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors,
assume any liability for any injury and/or damage to persons or property as a matter of
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products, instructions, or ideas contained in the material herein.
ISBN: 978-0-443-15307-5

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Publisher: Stacy Masucci

Acquisitions Editor: Linda Versteeg-Buschman
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 Dedication

This book is dedicated with gratitude to all the students

mentored during my tenure at the McGovern Medical
School, for their feedback and suggestions toward the
completion of this project. In addition, thanks goes to my
children, Jonas and Amanda, for sharing a thirst for
knowledge and for continuing to ask scientific questions;
and to my wife, Lori, for her love and patience and for
understanding my desire to complete this project. I also
wish to specially recognize my father, Paul Actor, PhD,
who instilled in me a sense of excitement about the
wonders of science and the curiosity to seek questions
about how biological systems function; and to my mother,
Ruthe Actor, who taught me to seek value in everything I
accomplish and to approach all challenges with an open
mind. Finally, I extend gratitude to Linda Versteeg-
Buschman at Elsevier for her continued support and
encouragement on this endeavor.

Jeffrey K. Actor
Contents

Preface xiii
Acknowledgments xvii

1. A functional overview of the immune system and

components 1
Chapter focus 1
Immune homeostasis 1
Self versus nonself 1
Innate and adaptive immunity 2
Anatomy of the immune system 4
Cells of the immune system 10
First-line defenders: the myeloid cells 12
Adaptive avengers: the lymphoid cells 14
Lymphocytes 14
Cluster of differentiation 16
Summary 18

2. The inflammatory response 21

Chapter focus 21
Inflammation 21
Initiation of the inflammatory response 22
The role of antibodies in inflammation 25
Biological functions of complement 25
Activation and directed migration of leukocytes 29
Pathogen recognition and cytokine signaling 30
Feedback and adaptation from a distance 31
Pathological consequences of the inflammatory Response 34
Summary 37

3. The B lymphocyte: antibodies and how they function 39

Chapter focus 39
B lymphocytes produce antibodies 39
Structural characteristics of immunoglobulins 39
Immunoglobulin nomenclature 43
Biologic properties of antibody isotypes 44
IgD 44

vii
viii Contents

IgM 44
IgG 45
IgA 46
IgE 47
Kinetics of antibody response 47
Membrane-bound immunoglobulin 48
Development of B cells 48
Antibody gene recombination 48
Development and selection of mature B cells 50
Activation and differentiation of B cells 51
Summary 52

4. T lymphocytes: ringleaders of adaptive

immune function 55
Chapter focus 55
T lymphocytes: specific and long-lasting immunity 55
The T cell receptor 55
T-cell development 56
Antigen recognition by T cells: requirement of major
histocompatibility molecules 59
The HLA locus 59
MHC class I 60
MHC class II 62
T lymphocyte functions 64
CD4 1 T-helper cells 66
Events involved in T lymphocyte activation 67
Role of T cells in B-cell activation 70
Cytotoxic T-cell effectors 71
CD8 1 T cells 71
Other lymphocyte phenotypes 73
γδ T cells 73
Innate lymphocytes 73
Natural killer T (NKT) cells 74
Superantigens 74
Summary 74

5. How we defend against infectious agents 79

Chapter focus 79
Immune homeostasis and pathogenic organisms 79
Major immune defense mechanisms against pathogens 79
Physical barriers to infection 80
Anatomic barrier 80
Physiological barrier 81
Phagocytic and endocytic barriers 81
Inflammatory barriers 81
 Contents ix

Bacterial infections 82
Microbial motifs detected through pattern-recognition receptors 85
Mycobacterial infections 88
Viral infections 91
Human immunodeficiency virus 93
Parasitic infections (helminths) 94
Fungal infections 97
Evasion of immune response 97
Summary 99

6. Basic disorders of immune function 101

Chapter focus 101
Immunodeficiency disorders 101
Genetic basis for primary immunodeficiency 101
Innate deficiencies 102
Chronic granulomatous disease/myeloperoxidase deficiency 102
Leukocyte adhesion deficiency 103
Chediak Higashi syndrome 103
Complement disorders 103
Innate pattern-recognition receptor disorders 104
Adaptive immune disorders 104
X-linked agammaglobulinemia 105
Selective immunoglobulin A deficiency 106
Common variable immunodeficiency 106
Hyper immunoglobulin M disorders 106
DiGeorge syndrome 107
Wiskott Aldrich syndrome 107
Severe combined immunodeficiency 107
Treatment of immunodeficiency diseases 108
Immunodeficiency as a predisposition to disease 109
Human immunodeficiency virus and acquired immunodeficiency
syndrome 109
Summary 110

7. Autoimmunity: regulation of response to self 111

Chapter focus 111
Homeostasis, immune regulation, and autoimmunity 111
Tolerance to self 112
Etiology of autoimmune disease 114
Role of autoantibodies and self-reactive T lymphocytes in
autoimmune disorders 118
Laboratory tests for autoimmunity 120
Targeted therapeutics 121
Summary 122
x Contents

8. Immune hypersensitivities 123

Chapter focus 123
Hypersensitive disorders 123
Type I hypersensitivity: immunoglobulin E mediated immediate
hypersensitivity 124
Type II hypersensitivity: antibody-mediated cytotoxic
hypersensitivity 126
Type III hypersensitivity: immune complex-mediated
hypersensitivity 128
Type IV hypersensitivity: delayed-type (cell-mediated)
hypersensitivity 129
Alternative hypersensitivity classifications 131
Summary 131

9. Vaccines and immunotherapy 133

Chapter focus 133
Principles of vaccination 133
Basic concepts of protective immunization 133
Types of immunizations 136
Age and timing of immunizations 137
Vaccine types and components 141
Immunologic adjuvants 144
Passive immunization 144
Therapeutic uses of immunoglobulins 145
Other ways to modify immunity 146
Summary 146

10. Cancer immunology 147

Chapter focus 147
Understanding immune defenses against cancers 147
Tumor antigens 147
Effector mechanisms in tumor immunity 149
Natural killer cells and innate response to tumor cells 150
Adaptive response to tumor cells 150
Escape mechanisms of tumor elimination 152
Tumors of the immune system 153
Immunodiagnosis and immunotherapy 154
Summary 156

11. Transplantation immunology 159

Chapter focus 159
Transplantation defined 159
Tissue histocompatibility 160
Natural isohemagglutinins 160
 Contents xi

Human leukocyte antigens 161

Allograft rejection 162
Hyperacute rejection 163
Accelerated rejection 163
Acute rejection 163
Chronic rejection 164
Graft-versus-host disease 164
Pretransplantation histocompatibility evaluation 165
Immunosuppressive drugs to prevent allograft rejection 166
Immunosuppressive therapy 167
Summary 168

12. Assessment of immune parameters and

immunodiagnostics 169
Chapter focus 169
Antibody antigen reactions 169
Affinity 169
Secondary manifestations of antibody antigen binding 170
Solid-phase precipitation assays 174
Radial immunodiffusion 174
The Ouchterlony assay 174
Immunoelectrophoresis 175
Latex agglutination 176
Lateral flow 176
Western blot 176
Immunoassays 177
Enzyme-linked immunoadsorbent assay 177
Detection of cellular antigens 178
Immunohistochemistry 180
Fluorescence-activated cell sorting analysis 180
Multiplex bead arrays 181
Assays to determine immune function 182
Complement fixation test 182
Phagocyte function assays 183
Lymphocyte function assays 185
Other tools to measure immunological status 187
Monoclonal antibodies 187
Microarrays 187
Summary 188

Glossary 189
Index 197
Preface

Our bodies have evolved a protective set of mechanisms, comprising cells

and organs, as a primary defense to maintain health. In essence, we have
developed internal tools to preserve health and homeostasis. Indeed, a work-
ing definition of health embraces the effective elimination or control of life-
threatening agents. This includes both infectious agents attacking from the
outside and internal threats, such as tumors. Immune responses are therefore
specifically designed to interact with, and respond to, the environment to
protect the host against pathogenic invaders and internal dangers. The goal
of this book is to appreciate the components of the human immune system
that work together to confer protection and prevent clinical disease.
We will begin our discussion by establishing a foundation for subsequent
chapters, through the presentation of the systems and cells involved in
immune responses. Chapter 1 will give a general overview of the mechan-
isms in place to fight against disease. Components and pathways will be
defined to allow the presentation of concepts of innate (always present) and
adaptive (inducible and specific) responses, and how these responses interact
with one another to form the basis for everyday protection. This edition
expands concepts of innate immune function, and phenotypic cellular sub-
sets, reflecting the discoveries that link innate and adaptive immunity. These
concepts will form the foundation to examine the process of defense against
various classes of pathogens. Chapter 2 will examine the coordinated effort
of cells and blood components in the development of inflammation as it is
related to protection against infection. Chapter 3 will introduce the basis for
the function of adaptive components, exploring the generation of B lympho-
cytes and the nature of antibodies. Chapter 4 will extend this discussion to
T-lymphocyte populations and examine how they serve as ringleaders for
immune function. Chapter 5 will discuss immune responses, with an element
of detail focused on commonly encountered infectious organisms. This over-
view also will include how the initial engagement of pathogens by innate
components triggers pathways that cause inflammation. A section of this
chapter will introduce the concept of opportunistic infections and diminished
response when individuals are immunocompromised.
Effective immune surveillance is paramount to maintaining health.
Chapter 6 will examine the basic disorders of immune function. Too little of
a response results in an inability to control threats, which thus is ineffective

xiii
xiv Preface

toward the elimination of infectious agents. This lack of reactivity (hyporeac-

tivity) leads to holes in our immune repertoire. This may be the result of
genetic deficiencies or acquired compromise of immune function. In the
same manner, responses representing excessive activity also can lead to dam-
age to the host. This overaggressive response, a state of hyperreactivity, may
reflect a productive response that increases in intensity and duration without
effective control. The dysregulation leads to tissue-damaging events and
eventual states of disease.
The chief function of the immune system is to distinguish between what
is you (self) and what constitutes external threats. When the ability to distin-
guish these elements is compromised, autoimmunity may arise. In Chapter 7,
autoimmune dysfunction will be addressed, moving from basic concepts to
the specific mechanisms involved in major clinical disorders. This includes a
detailed discussion of how the self is recognized, as well as the mechanisms
involved in tolerance that limit reactivity to our own tissues. The goals here
are to present the clinical manifestation of autoimmunity in such a manner
that outward symptoms are understood through investigation of the molecu-
lar targets involved in the host immune self-recognition response. At other
times, misdirected recognition of nonself elements, such as environmental
allergens that typically are considered harmless, results in the development
of clinical presentations. Therefore Chapter 8 will examine the processes
involved in the manifestation of immune dysfunction, including the concepts
of immune hypersensitivities that lead to clinical disease.
The general topic of vaccines will be addressed in Chapter 9, including
both how they work and a frank discussion of the relative truths surrounding
their use. Novel methodologies related to vaccine components and composi-
tion that are now globally adopted for clinical use will be detailed. This
chapter will also contain information on newly developed therapeutics that
are grounded in methods that lead to immune modification and factors that
promote a healthy immune response (e.g., lifestyle activities and good com-
mon practices). Indeed, it is critical that we maintain a healthy balance
throughout our lives to ensure functional immune response as we age. The
challenges faced at each stage of our lives, from that found in the prenatal/
newborn period, to midlife, to so-called maturity, are mentioned in a way
that encourages a healthy condition to allow the optimization of immune
function.
A discussion of natural (effective) response to tumor development in
Chapter 10 will lead to an investigation into the components of immune
function to eliminate potentially dangerous precancerous events naturally.
This will be followed by coverage of the challenges that we face when pro-
tective responses fail and tumors develop. A section also will contain infor-
mation on cancers of the immune system and the problems that arise when
the protective cells themselves become the cause of tumorigenic activity.
Advances in immune-based cancer therapies will be described that include
 Preface xv

subjects such as CAR-T therapy, directed monoclonal treatments, and path-

way inhibitors.
Chapter 11 will delve once more into details underlying the concepts of
“self” versus “nonself” and blood types, with the goal to present genetic rela-
tionships (similarities as well as differences) between individuals. The
mechanisms of the immunobiology of transplantation will be discussed, with
details about the contributing cells and factors involved in transplant accep-
tance and rejection. The challenge is to appreciate the importance of innate
and adaptive components in graft recognition, as well as to recognize the
clinical consequences of transplantation that affect aspects of daily activities.
Rejection topics will be discussed, including graft-versus-host disease, as
well as modern immune-based therapeutics designed to alter immune func-
tion to limit graft rejection.
Finally, additional information and resources will be provided in
Chapter 12 to allow readers to develop an immune-based foundation of
knowledge to understand the clinical tests associated with identifying
immune parameters that arise during the development of disease states. As
such, this includes an introduction to mechanisms that form the basis of
immune-related diagnostics and identification of immune properties of the
blood during disorders and how immune factors influence outcomes of dis-
ease states.
All in all, the hope is that this book will present the concept of the
immune system in a holistic manner so that readers may better understand
immune-based diseases resulting from either immune system component
deficiencies or excess activity. This book is aimed at those who want to
know more to encourage readers to explore deeper. It is aimed at the curious
who have never previously considered the underlying facets of effective
immune function, students who wish to expand their basic knowledge of bio-
logical systems, physicians seeking to refresh their understanding of the
immune concepts that cause clinical disease, nurses who desire to expand
their view of symptom development in patients, patients who want a simple
explanation for the complex way that their bodies respond in the context of
the world they live in, and, finally, all who seek to find out how the body
confers protection against infectious agents, maintain everyday homeostasis,
and guard against dysregulation of the normal response to confer health and
control the development of disease.
Acknowledgments

I would like to give special credit to Keri C. Smith, PhD, for academic con-
tribution to the chapters on inflammation, antibodies, and immunoassays;
and to Robert L. Hunter, MD, PhD, for sharing his positive outlook on life
and showing me the joy of embracing scientific thought. In addition, I would
like to acknowledge Jonas A. Actor, PhD; Jun Liu, PhD; and Angel Paredes,
PhD, for the front cover 3D scanning electron micrograph of Mycobacterium
tuberculosis infected macrophages.

xvii
Chapter 1

A functional overview of the

immune system and
components
Chapter focus
To establish a foundation to appreciate how components of the immune sys-
tem work together to protect against the development of clinical disease. The
basic systems and cells involved in immune responses will be presented in
this chapter to give a general overview of functional immunity. Components
and systems will be defined to allow an understanding of concepts of innate
(always present) and adaptive (inducible and specific) responses, and how
these responses interact with one another to form the basis for protection
against disease.

Immune homeostasis
A functional immune system offers constant surveillance of human beings in
relationship to the world. It confers a balanced state of health through effec-
tive elimination of infectious agents (bacteria, viruses, fungi, and parasites)
and through control of malignancies. Indeed, the immune system has evolved
to allow cells and organs to interact with the environment to protect against
harmful invaders. At the same time, mechanisms are in place to instill toler-
ance toward the naturally occurring microbiome (i.e., microbial and viral
agents) that reside within us in symbiotic ways. Taken together, these
responses represent a balance of components that ward off the development
of clinical disease.

Self versus nonself

Discrimination between the self and the nonself is considered the chief func-
tion of the immune system. We are under constant assault by invaders. Our
bodies represent prime substrates for organisms to grow and reside, with an
abundance of nutrients, warmth, and protection from the outside elements.
The immune system is basically a series of obstacles to limit and inhibit

Introductory Immunology. DOI: https://doi.org/10.1016/B978-0-443-15307-5.00004-4

© 2023 Elsevier Inc. All rights reserved. 1
2 Introductory Immunology

pathogen entry and then to attack and destroy those organisms once they
enter the body. The immune response is exquisitely designed to recognize
these invaders as “foreign.” In fact, the major feature that renders our
immune system so effective is its ability to distinguish our body’s own cells
(the self) from that which it considers foreign (termed the nonself). Each of
our cells carries specific tags, or molecular markers, that label it as “self.”
These markers are important, as they not only determine what is unique
about us but also distinguish one person from another.
Almost anything and everything that registers as “nonself” will trigger an
immune response. An intricate system of molecular communication and cellu-
lar interactions allows immune components to function in concert to combat
disease-causing organisms. The foreign agent (microbe, virus, parasite, etc.), or
any part of it that can be specifically recognized, is called an antigen. Simply
put, an antigen is defined as any substance or physical structure that can be
recognized by the immune system. Major classes of antigens include proteins,
carbohydrates, lipids, and nucleic acids. If an antigen is of high complexity and
weight, it can trigger full immune activity and become immunogenic.
The ability to distinguish our own cells from the outside world is critical
to maintain functional protection. If this ability is lost (e.g., when “self” tis-
sue is seen as foreign), then our immune system launches an aggressive
response against our own tissues. This is what happens during autoimmu-
nity, where destruction of the self leads to clinical disease.
The immune system maintains a balance of responsiveness. Too little a
response is ineffective, while too aggressive a response can lead to the tar-
geted destruction of bystander tissues. Both scenarios are equally as devastat-
ing and may result in clinical disease. The regulation of immune function
and overall immuno-homeostasis is under the control of multiple factors,
including genetic components and environmental cues. The intensity and
duration of the responses must be sufficient to protect against invading
pathogens, with prompt and specific downregulation when the foreign mate-
rial (the antigen or the pathogen) is no longer present. The clinical state that
arises when immune responses are not properly regulated is termed hyper-
sensitivity, a state of excessive or inappropriate responses leading to disease.
As one might imagine, hypersensitivity can occur in many different forms,
depending upon which arm of the immune system is dysregulated.

Innate and adaptive immunity

The immune system is loosely divided into two major functional categories
termed innate and adaptive immunity. Innate immune mechanisms provide
the first line of defense from infectious diseases (Table 1.1). The innate
immune components exist from birth and consist of components available
prior to the onset of infection. These defensive components include both
physical barriers and biochemical factors. Defensive innate mechanisms may
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