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 ENCYCLOPEDIA OF

INFECTION
AND IMMUNITY
VOLUME

\
Editor in Chief
_
ELSEVIER Nima Rezaei
 ENCYCLOPEDIA OF
INFECTION AND IMMUNITY
 ENCYCLOPEDIA OF
INFECTION AND IMMUNITY
EDITOR-IN-CHIEF
Nima Rezaei
Research Center for Immunodeficiencies, Children’s Medical Center,
Tehran University of Medical Sciences, Tehran, Iran

Department of Immunology, School of Medicine,

Tehran University of Medical Sciences, Tehran, Iran

Network of Immunity in Infection, Malignancy and Autoimmunity (NIIMA),

Universal Scientific Education and Research Network (USERN), Stockholm, Sweden

VOLUME 1

Section Editors
Section 1: The Immune System, Edited by Nima Rezaei
Section 2: Pathogens and Immunity, Edited by Rowena Jenkins
Section 3: Bacteria, Edited by Fernando Cobo
Elsevier
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EDITOR-IN-CHIEF

Nima Rezaei gained his medical degree (MD) from Tehran University of
Medical Sciences and subsequently obtained an MSc in molecular and genetic
medicine and a PhD in clinical immunology and human genetics from the
University of Sheffield, UK. He also spent a short-term fellowship of pediatric
clinical immunology and bone marrow transplantation in the Newcastle General
Hospital. Professor Rezaei is now the full professor of immunology and vice dean
of International Affairs, School of Medicine, Tehran University of Medical Sci-
ences, and the co-founder and head of the Research Center for Immunodefi-
ciencies. He is also the founding president of Universal Scientific Education and
Research Network (USERN). Prof. Rezaei has already been the director of more
than 100 research projects and has designed and participated in several interna-
tional collaborative projects. Prof. Rezaei is the editor, editorial assistant, or
editorial board member of more than 40 international journals. He has edited
more than 50 international books, has presented more than 500 lectures/posters
in congresses/meetings, and has published more than 1100 scientific papers in
the international journals.

v
EDITORIAL BOARD

Editor-in-Chief

Nima Rezaei
Research Center for Immunodeficiencies, Children’s Medical Center, Tehran University of Medical Sciences, Tehran, Iran;
Department of Immunology, School of Medicine, Tehran University of Medical Sciences, Tehran, Iran; Network of
Immunity in Infection, Malignancy and Autoimmunity (NIIMA), Universal Scientific Education and Research Network
(USERN), Stockholm, Sweden

Section Editors

Section 1. The Immune System

Nima Rezaei
Research Center for Immunodeficiencies, Children’s Medical Center, Tehran University of Medical Sciences, Tehran, Iran;
Department of Immunology, School of Medicine, Tehran University of Medical Sciences, Tehran, Iran; Network of
Immunity in Infection, Malignancy and Autoimmunity (NIIMA), Universal Scientific Education and Research Network
(USERN), Stockholm, Sweden

Section 2. Pathogens and Immunity

Dr Rowena Jenkins
Swansea University Medical School, Swansea, United Kingdom

Section 3. Bacteria
Fernando Cobo
Department of Microbiology and Instituto de Investigación Biosanitaria ibs.GRANADA, University Hospital Virgen de las
Nieves, Granada, Spain

Section 4. Viruses
Nima Rezaei
Research Center for Immunodeficiencies, Children’s Medical Center, Tehran University of Medical Sciences, Tehran, Iran;
Department of Immunology, School of Medicine, Tehran University of Medical Sciences, Tehran, Iran; Network of
Immunity in Infection, Malignancy and Autoimmunity (NIIMA), Universal Scientific Education and Research Network
(USERN), Stockholm, Sweden

Section 5. Fungi, Protozoan Parasites, and Helminths

Fungi
Fanny Lanternier
Hôpital Universitaire Necker-Enfants malades, Assistance Publique—Hôpitaux de Paris, Paris, France; Université de
Paris, Institut Pasteur, Unité de Mycologie Moléculaire, UMR 2000, CNRS, CNR des Mycoses Invasives et antifongiques,
Paris, France

Protozoan Parasites and Helminths

Fabrizio Bruschi, MD
Department of Translational Research, N.T.M.S., Università di Pisa, Pisa, Italy

vii
viii Editorial Board

Section 6. Insect and Mites

Prof. Giovanni Benelli, Ph.D.
Department of Agriculture, Food and Environment, University of Pisa, Pisa, Italy

Section 7. Clinical Infectious Disease

Federico Iovino, PhD.
Associate Professor and Principal Investigator, Iovino Group, Laboratory of Neuro-Infections and Neuro-Inflammation,
Department of Neuroscience – Karolinska Institutet, Stockholm, Sweden

Section 8. Clinical Immunology

Hans D. Ochs
University of Washington and Seattle Children’s Research Institute, Seattle, WA, United States

Nima Rezaei
Research Center for Immunodeficiencies, Children’s Medical Center, Tehran University of Medical Sciences, Tehran, Iran;
Department of Immunology, School of Medicine, Tehran University of Medical Sciences, Tehran, Iran; Network of
Immunity in Infection, Malignancy and Autoimmunity (NIIMA), Universal Scientific Education and Research Network
(USERN), Stockholm, Sweden

Section 9. Laboratory Tests

Nima Rezaei
Research Center for Immunodeficiencies, Children’s Medical Center, Tehran University of Medical Sciences, Tehran, Iran;
Department of Immunology, School of Medicine, Tehran University of Medical Sciences, Tehran, Iran; Network of
Immunity in Infection, Malignancy and Autoimmunity (NIIMA), Universal Scientific Education and Research Network
(USERN), Stockholm, Sweden

Section 10. Treatment Protocols

Nima Rezaei
Research Center for Immunodeficiencies, Children’s Medical Center, Tehran University of Medical Sciences, Tehran, Iran;
Department of Immunology, School of Medicine, Tehran University of Medical Sciences, Tehran, Iran; Network of
Immunity in Infection, Malignancy and Autoimmunity (NIIMA), Universal Scientific Education and Research Network
(USERN), Stockholm, Sweden
LIST OF CONTRIBUTORS FOR VOLUME 1

Raquel Abad Jorge Arca-Suarez

National Reference Laboratory for Neisserias, National Department of Microbiology, Complexo Hospitalario
Centre of Microbiology—Instituto de Salud Carlos III, Universitario A Coruña (CHUAC), Instituto de
Majadahonda, Spain Investigación Biomédica A Coruña (INIBIC), A Coruña,
Spain
Hassan Abolhassani
Division of Clinical Immunology, Department of Beatriz Plata Barril
Laboratory Medicine, Karolinska Institute at Karolinska Department of Infectious Diseases and Clinical
University Hospital Huddinge, Stockholm, Sweden Microbiology, Jaén University Hospital, Jaén, Spain
Jeffrey K Actor Jay A Berzofsky
University of Texas Health Science Center, McGovern Vaccine Branch, Center for Cancer Research, National
Medical School, Department of Pathology and Laboratory Cancer Institute, NIH, Bethesda, MD, United States
Medicine, Houston, TX, United States
Jaime Borrego
Jawad Ahmed Hospital Universitario Virgen de las Nieves. Granada, Spain
Department of Internal Medicine, Dow University of
Helen Louise Brown
Health Sciences, Karachi, Pakistan
School of Dentistry, Cardiff University, Cardiff, United
Mahzad Akbarpour Kingdom
Network of Immunity in Infection, Malignancy and
Elizabeth Calatrava
Autoimmunity (NIIMA), Universal Scientific Education
Department of Microbiology and Instituto de Investigación
and Research Network (USERN), Tehran, Iran;
Immunology Board for Transplantation and Cell-Based Biosanitaria, ibs. GRANADA, University Hospital Virgen
de las Nieves, Granada, Spain
Therapeutics (ImmunoTACT), Universal Scientific
Education and Research Network (USERN), Tehran, Iran; Noelia Calvo Sánchez
Advanced Cellular Therapeutics Facility (ACTF), Servicio de Microbiologí a, Hospital Clí nico Universitario de
Hematopoietic Cellular Therapy Program, Section of Salamanca, Salamanca, Spain; Departamento de Ciencias
Hematology & Oncology, Department of Medicine, Biomédicas y del Diagnóstico. Facultad de Medicina,
University of Chicago Medical Center, Chicago, United Universidad de Salamanca, Salamanca, Spain
States
Isabel Casanovas Moreno-Torres
Luis Aliaga Department of Microbiology, Hospital Santa Ana,
Department of Medicine, University of Granada, Granada, Granada, Spain
Spain; Department of Internal Medicine, Hospital
Universitario San Cecilio, Granada, Spain Fernando Cobo
Department of Microbiology and Instituto de Investigación
Jorge Amich
Biosanitaria ibs.GRANADA, University Hospital Virgen de
Manchester Fungal Infection Group (MFIG), Division
las Nieves, Granada, Spain
of Infection, Immunity and Respiratory Medicine,
School of Biological Sciences, Faculty of Biology, Claudio Cortes
Medicine and Health, University of Manchester, Department of Foundational Medical Studies, Oakland
Manchester Academic Health Science Centre, University William Beaumont School of Medicine,
Manchester, United Kingdom Rochester, MI, United States

ix
x List of Contributors for Volume 1

Elena Cuadros Moronta Esther Gómez-Vicente

Department of Clinical Microbiology, Sierrallana Hospital, Department of Microbiology, Hospital Virgen de las Nieves,
Torrelavega, Spain Granada, Spain
Francisco de Asís Ramirez Laura Viñuela González
Primary Care Center “El Valle", Jaén, Spain; Centro de Microbiology Department, Hospital Universitario Clí nico
Salud “El Valle”, Jaén, Spain San Cecilio, Granada, Spain
Adolfo de Salazar González A González-Martínez
Department of Microbiology, Hospital Universitario Clí nico Servicio de Microbiologí a, Hospital Universitario Virgen de
San Cecilio, Granada, Spain las Nieves, Granada, Spain
Joshua De Sousa Casal Juan Francisco Gutiérrez Bautista
Department of Immunology, University of Toronto, Hospital Universitario Virgen de las Nieves, Servicio de
Toronto, ON, Canada; Sunnybrook Research Institute, Análisis Clí nicos e Inmunologí a, Granada, Spain
Toronto, ON, Canada
Sofia Hain
Zlatko Dembic Institute of Immunology and Immunotherapy, University of
Molecular Genetics Laboratory, Department of Oral Biology, Birmingham, Birmingham, United Kingdom
Faculty of Dentistry, University of Oslo, Oslo, Norway
Llinos G Harris
Roman Deniskin
Microbiology and Infectious Diseases, Institute of Life
University of Kentucky and Kentucky Children's Hospital, Sciences, Medical School, Faculty of Medicine, Health and
Lexington, KY, United States
Life Science, Swansea University, Swansea, United
Rebecca A Drummond Kingdom
Institute of Immunology and Immunotherapy, University of Jenny A Herbert
Birmingham, Birmingham, United Kingdom; Institute of
University of Manchester, Manchester, United Kingdom
Microbiology and Infection, University of Birmingham,
Birmingham, United Kingdom Paula Hernández Calvo
Servicio de Microbiologí a, Hospital Clí nico
Noha Mousaad Elemam Universitario de Salamanca, Salamanca, Spain;
Department of Clinical Sciences, College of Medicine,
Departamento de Ciencias Biomédicas y del Diagnóstico.
University of Sharjah, Sharjah, United Arab Emirates;
Facultad de Medicina, Universidad de Salamanca,
Immuno-Oncology Group, Sharjah Institute for Medical
Salamanca, Spain
Research (SIMR), Sharjah, United Arab Emirates
Savannah B Howe
Pooya Farhangnia
Vaccine Branch, Center for Cancer Research, National
Department of Immunology, School of Medicine, Iran
Cancer Institute, NIH, Bethesda, MD, United States
University of Medical Sciences, Tehran, Iran; Network of
Immunity in Infection, Malignancy and Autoimmunity Imogen Anne Jones
(NIIMA), Universal Scientific Education and Research Faculty of Health, School of Biomedical Sciences,
Network (USERN), Tehran, Iran; Immunology Board for University of Plymouth, Plymouth, United Kingdom
Transplantation and Cell-Based Therapeutics
Lovleen Tina Joshi
(ImmunoTACT), Universal Scientific Education and
Faculty of Health, School of Biomedical Sciences,
Research Network (USERN), Tehran, Iran
University of Plymouth, Plymouth, United Kingdom
Viviana P Ferreira
Mahdis Keshavarz-Fathi
Department of Medical Microbiology and Immunology,
University of Toledo College of Medicine and Life Sciences, Cancer Immunology Project (CIP), Universal Scientific
Education and Research Network (USERN), Tehran, Iran;
Toledo, OH, United States
School of Pharmacy, Shahid Beheshti University of Medical
Carla Foronda García-Hidalgo Sciences, Tehran, Iran
Hospital Universitario Virgen de las Nieves, Granada,
Spain Mahsa Keshavarz-Fathi
School of Medicine, Tehran University of Medical Sciences,
Cristina Gómez-Camarasa Tehran, Iran; Cancer Immunology Project (CIP), Universal
Microbiology Department, Hospital Universitario Clí nico Scientific Education and Research Network (USERN),
San Cecilio, Granada, Spain Tehran, Iran
 List of Contributors for Volume 1 xi

Bariaa A Khalil Pavel P Nesmiyanov

Department of Clinical Sciences, College of Medicine, Medical Research and Education Center, Lomonosov
University of Sharjah, Sharjah, United Arab Emirates; Moscow State University (Moscow State University
Immuno-Oncology Group, Sharjah Institute for Medical Clinic), Moscow, Russia; Engelhardt Institute of
Research (SIMR), Sharjah, United Arab Emirates Molecular Biology, Russian Academy of Sciences, Moscow,
Russia
Ana Lara-Oya
Department of Microbiology, Hospital Universitario de MJ Olivares-Duran
Jaén, Jaén, España Servicio de Análisis Clí nicos e Inmunologí a, Hospital
Universitario Virgen de las Nieves, Granada, Spain
Carmen Liébana-Martos
Facultativo Especialista de Área en Microbiologí a, Servicio Purevdorj B Olkhanud
de Microbiologí a, Hospital Universitario de Jáen, Jaén, Vaccine Branch, Center for Cancer Research, National
España Cancer Institute, NIH, Bethesda, MD, United States

Ana Fuentes López Anastasia Sampedro Padilla

Microbiology Department, Hospital Universitario Clí nico Hospital Neurotraumatológico del HU de Jaén, Jaén, Spain
San Cecilio, Granada, Spain Stavros Panagiotou
Azzam A Maghazachi University of Manchester, Manchester, United Kingdom
Department of Clinical Sciences, College of Medicine, Samira Rajaei
University of Sharjah, Sharjah, United Arab Emirates; Immunology Department, School of Medicine, Tehran
Immuno-Oncology Group, Sharjah Institute for University of Medical Sciences, Tehran, Iran
Medical Research (SIMR), Sharjah, United Arab
Emirates JA Reguera Márquez
Servicio de Microbiologí a, Hospital Universitario Virgen de
Farheen Malik las Nieves, Granada, Spain
Department of Internal Medicine, Dow University of
Nima Rezaei
Health Sciences, Karachi, Pakistan
Department of Immunology, School of Medicine, Tehran
Josefa Martínez University of Medical Sciences, Tehran, Iran; Network of
Department of Internal Medicine, Hospital Universitario Immunity in Infection, Malignancy and Autoimmunity
San Cecilio, Granada, Spain (NIIMA), Universal Scientific Education and Research
Network (USERN), Tehran, Iran; Research Center for
Hamid Reza Mirzaei
Immunodeficiencies, Children's Medical Center, Tehran
Department of Medical Immunology, School of Medicine,
University of Medical Sciences, Tehran, Iran; Cancer
Tehran University of Medical Sciences, Tehran, Iran
Immunology Project (CIP), Universal Scientific
Hamid-Reza Mohammadi-Motlagh Education and Research Network (USERN), Tehran,
Medical Biology Research Center, Health Technology Iran
Institute, Kermanshah University of Medical Sciences,
Johanna Rhodes
Kermanshah, Iran
MRC Centre for Global Disease Analysis, Imperial College
Sara Momtazmanesh London, London, United Kingdom
Research Center for Immunodeficiencies, Children's
Lothar Rink
Medical Center, Tehran University of Medical Sciences,
Institute of Immunology, Medical Faculty, RWTH Aachen
Tehran, Iran; Network of Immunity in Infection,
University, Aachen, Germany
Malignancy and Autoimmunity (NIIMA), Universal
Scientific Education and Research Network (USERN), Javier Rodríguez-Granger
Tehran, Iran Department of Microbiology, Hospital Universitario Virgen
de las Nieves, Granada, Spain
Manuela Moreno-Higueras
Department of Internal Medicine, Hospital Universitario Laura Lucía Rojas-García
San Cecilio, Granada, Spain Department of Microbiology, University Hospital San
Cecilio, Granada, Spain
Juan Luis Muñoz Bellido
Servicio de Microbiologí a, Hospital Clí nico Universitario de Carlos Rosales
Salamanca, Salamanca, Spain; Departamento de Ciencias Departamento de Inmunologí a, Instituto de Investigaciones
Biomédicas y del Diagnóstico. Facultad de Medicina, Biomédicas, Universidad Nacional Autónoma de México,
Universidad de Salamanca, Salamanca, Spain Mexico City, Mexico
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xii List of Contributors for Volume 1

Mona Sadeghalvad Eileen Uribe-Querol

Department of Immunology, School of Medicine, Tehran División de Estudios de Posgrado e Investigación, Facultad
University of Medical Sciences, Tehran, Iran; Network of de Odontologí a, Universidad Nacional Autónoma de
Immunity in Infection, Malignancy and Autoimmunity México, Mexico City, Mexico
(NIIMA), Universal Scientific Education and Research
Sylvia Valdezate
Network (USERN), Tehran, Iran; Research Center for
Reference and Research Laboratory for Taxonomy,
Immunodeficiencies, Children's Medical Center, Tehran
National Centre of Microbiology, Instituto de Salud Carlos
University of Medical Sciences, Tehran, Iran
III, Madrid, Spain
Antonio Sampedro
Julio A Vázquez
Microbiology Service, Virgen de las Nieves University
National Reference Laboratory for Neisserias, National
Hospital, Granada, Spain
Centre of Microbiology—Instituto de Salud Carlos III,
Esther Serrano-Conde Sánchez Majadahonda, Spain
Microbiology Department, Hospital Universitario Clí nico
Inga Wessels
San Cecilio, Granada, Spain
Institute of Immunology, Medical Faculty, RWTH Aachen
Lisa Forbes Satter University, Aachen, Germany
Baylor College of Medicine and Texas Children's Hospital, Rachael C Wilkinson
Houston, TX, United States Sir William Dunn School of Pathology, Oxford, United
Jennifer Scott Kingdom
Manchester Fungal Infection Group (MFIG), Division Thomas S Wilkinson
of Infection, Immunity and Respiratory Medicine, Institute of Life Science, Microbiology and Infectious
School of Biological Sciences, Faculty of Biology, Disease, Swansea University Medical School (SUMS),
Medicine and Health, University of Manchester, Swansea, United Kingdom
Manchester Academic Health Science Centre,
Manchester, United Kingdom Catrin F Williams
School of Biosciences, Cardiff University, Cardiff, Wales,
Katie Silver United Kingdom
Faculty of Health, School of Biomedical Sciences,
University of Plymouth, Plymouth, United Kingdom Lisa K Williams
Department of Agriculture and Animal Science, Hartpury
Keri C Smith University, Gloucester, United Kingdom
SABA Medical School, SABA, Dutch Caribbean
Juan Carlos Zúñiga-Pflücker
Gabriele Sorci Department of Immunology, University of Toronto,
Biogéosciences, CNRS UMR 6282, Université de Toronto, ON, Canada; Sunnybrook Research Institute,
Bourgogne Franche-Comté, Dijon, France Toronto, ON, Canada
FOREWORD

It is a great honor to be asked to introduce, with this foreword, the Encyclopedia of Infection and Immunity. This
work will be read and enjoyed by many students, both graduate and postgraduate, including clinicians,
laboratory practitioners, and scientists. Many colleagues have contributed to this important work, with over
230 original chapters assembled in sections and listed alphabetically. I offer my warm congratulations to the
editors of this book, for soliciting a wide body of expertise and assembling it in a form that gives so much
pleasure and information to the reader.
For those of us who work in the field of infection and immunity, the last two years have been exceptionally
challenging. At the current time we are 22 months into a major pandemic caused by a novel coronavirus and its
variants, and worldwide, sadly, there have been many hospitalizations and deaths. Clinicians and scientists in
our field have been working flat out, caring for patients, generating or utilizing diagnostic tests, generating new
knowledge in immunity to coronaviruses, or discovering, testing, and deploying new vaccines. Public Health
practitioners have organized our medico-social responses to limit transmission and deal with the consequences
of infection. Behavioral scientists have transformed our approach to informing the public, getting the best
possible outcomes from effective communication. Governments have learned the danger of ignoring the
expertise of scientists, and in many countries, the public look upon clinicians and scientists with gratitude
and respect. It is a testament to the advancement of infection and immunity science that the genome of the virus
was sequenced within a few weeks of COVID-19 emerging, and that within 12 months, the first trials of novel
vaccines had been completed. These vaccines included some with exceptionally novel technologies, including
the RNA vaccines and the viral-vectored vaccines.
Underpinning the academic agility of students and practitioners in Infection and Immunity is the universal
strength of sound education. Readers of this book will be seeking the knowledge they need for a comprehensive
understanding of infectious disease and immunology. We work in a fascinating discipline. Our field is unique in
that pathogenesis involves at least two sets of genomes—on the one hand the microbe, and on the other, the
host. The microbial genome programs its microenvironment to enable acquisition of and residence in a niche
for sufficient time to replicate and further disseminate to other hosts. The human genome has evolved to
tolerate and contain a wide range of symbionts and pathobionts, with highly sophisticated immune
decision-making at the cellular level, but to respond to invading species with an elaborate coordinated
repertoire of innate and acquired immunity. The complex conversation between many different microbes and
the human host generates a wide range of disease phenotypes. This makes the job of an infectious disease
physician so challenging but at the same time so fulfilling.
Our field is enriched by the many scientists and clinicians who are willing to share their experience with their
peers and students. This book reflects the great variety of knowledge that has emerged over centuries of inquiry
and endeavor, and I believe you will be rewarded by using it as one of your reference sources.

Robert C. Read
Editor-in-Chief, Journal of Infection
Professor of Infectious Diseases and Consultant Physician
University Hospital Southampton, United Kingdom

xiii
PREFACE

Although infections have been one of the most important health threats since centuries ago, the mechanisms of
human defense against infections have been unclear until last century. The scientific world has remarkable
progresses in the field of immunology during recent decades. Indeed, the novel discovery of genes related to
different pathogens has enhanced our knowledge about the immune system. Understanding different patho-
gens and the immune mechanisms promote diagnostic strategies and design more efficient therapeutic agents.
Encyclopedia of Infection and Immunity is a comprehensive text, which covers topics not only on basic
immunology and microbiology but also on clinical immunology and infectious diseases. Section 1 focuses
on the immune system, while Section 2 describes the interaction between pathogens and immunity. Bacteria,
viruses, fungi, protozoan parasites and helminths, and insect and mites are explained in Sections 3–6, respec-
tively. Section 7 discusses the clinical infectious disease challenges, whereas clinical immunology is covered in
Section 8. Laboratory tests and treatment protocols are the main focuses of Sections 9 and 10, respectively.
Encyclopedia of Infection and Immunity is the result of valuable contribution of more than 400 scientists from
many well-known universities/institutes worldwide. I would like to hereby acknowledge the expertise of all
contributors, for generously devoting their time and considerable effort in preparing their respective chapters.
I would also like to express my gratitude to Elsevier for providing us the opportunity to publish this
Encyclopedia.
Finally, I hope that this comprehensive book will be of special value for researchers and clinicians who wish
to extend their knowledge on infection and immunity.

Nima Rezaei, MD, PhD

xv
CONTENTS OF VOLUME 1

Section 1: The Immune System, Edited by Nima Rezaei

General Concepts of Immunity 1
Nima Rezaei, Mona Sadeghalvad, and Hamid-Reza Mohammadi-Motlagh

Organs and Tissues of the Immune System 14

Hamid-Reza Mohammadi-Motlagh, Mona Sadeghalvad, and Nima Rezaei

Structure and Function of the Immune System 24

Mona Sadeghalvad, Hamid-Reza Mohammadi-Motlagh, and Nima Rezaei

Adaptive Immunity 39
Hamid Reza Mirzaei

Cell-Mediated Immunity 56
Samira Rajaei

T Cell Development 64
Joshua DeSousa Casal and Juan Carlos Zúñiga-Pflücker

Antigens 76
Jay A Berzofsky, Savannah B Howe, and Purevdorj B Olkhanud

Antigen Presentation and Major Histocompatibility Complex 90

Pavel P Nesmiyanov

Phagocytosis 99
Eileen Uribe-Querol and Carlos Rosales

Dendritic Cells 110

Pavel P Nesmiyanov

Natural Kills Cells 118

Roman Deniskin and Lisa Forbes Satter

Toll-Like Receptors 130

Mona Sadeghalvad, Hamid-Reza Mohammadi-Motlagh, and Nima Rezaei

The Complement System 144

Viviana P Ferreira and Claudio Cortes

Cytokines in Innate Immunity 170

Zlatko Dembic

Chemokines and Chemokine Receptors 193

Noha Mousaad Elemam, Bariaa A Khalil, and Azzam A Maghazachi

Immunological Tolerance 206

Pooya Farhangnia and Mahzad Akbarpour

Immunologic Memory 221

Hassan Abolhassani

xvii
xviii Contents of Volume 1

Inflammation 230
Jeffrey K Actor and Keri C Smith

Hypersensitivity 243
Sara Momtazmanesh and Nima Rezaei

Immunosenescence 259
Lothar Rink and Inga Wessels

Cancer Immunology 277

Mahsa Keshavarz-Fathi, Mahdis Keshavarz-Fathi, and Nima Rezaei

Section 2: Pathogens and Immunity, Edited by Rowena Jenkins

Normal Microbial Flora 297
Lovleen Tina Joshi, Imogen Anne Jones, and Katie Silver

Microbial Diversity and Classification 306

Rachael C Wilkinson

Microbial Growth 324

Helen Louise Brown

Microbial Infections: The Good, the Bad and the Ugly 336
Lisa K Williams

Microbial Cell Structure and Organization: Bacteria 345

Llinos G Harris

Microbial Metabolism 363

Catrin F Williams

Primary Metabolism of Human Pathogenic Fungi, Importance for Virulence and Potential for Drug
Development 377
Jennifer Scott and Jorge Amich

Immunity to Bacterial Infections 408

Thomas S Wilkinson

Immune Response to Viruses 429

Jenny A Herbert and Stavros Panagiotou

Immunity to Fungal Infections 445

Sofia Hain and Rebecca A Drummond

Microbial Genetics in Mycology 462

Johanna Rhodes

Immunology of Parasitism 474

Gabriele Sorci

Section 3: Bacteria, Edited by Fernando Cobo

General Characteristics of Bacteria 481
Paula Hernández Calvo, Noelia Calvo Sánchez, and Juan Luis Muñoz Bellido

Pathogenesis of Bacterial Infections and Bacterial Persistence 492

Fernando Cobo

Staphylococcus and Other Catalase-Positive Cocci 498

Carla Foronda Garcí a-Hidalgo

Streptococcus pneumoniae 511

Jawad Ahmed and Farheen Malik

Other Streptococcus Species and Enterococcus 529

Elizabeth Calatrava
 Contents of Volume 1 xix

Genus Neisseria 542

Raquel Abad and Julio A Vázquez

Corynebacterium diphtheriae 559

Jaime Borrego

Listeria and Erysipelothrix 573

Cristina Gómez-Camarasa

Coryneform Gram-Positive Bacilli 580

Laura Lucí a Rojas-Garcí a

Nocardia, Rhodococcus, Streptomyces and Other Aerobic Actinomycetes 589

Sylvia Valdezate

Infections Caused by Anaerobic Microorganisms 614

Fernando Cobo

Human Pathogenic Enterobacterales 628

Laura Viñuela González

Vibrio 637
Ana Fuentes López, Esther Serrano-Conde Sánchez, Laura Viñuela González, and Cristina Gómez Camarasa

Pseudomonas Infections 644

Luis Aliaga, Manuela Moreno-Higueras, Josefa Martí nez, and Javier Rodrí guez-Granger

Stenotrophomonas, Burkholderia and Other Related Microorganisms 656

Ana Lara-Oya

Moraxella and Other Non-Fermentative Gram-Negative Bacilli 662

Esther Gómez-Vicente

Francisella, Brucella and Pasteurella 673

Beatriz Plata Barril

Campylobacter and Helicobacter 685

Isabel Casanovas Moreno-Torres

Haemophilus, Bordetella and Bartonella 694

Adolfo de Salazar González and Jorge Arca-Suarez

Mycobacterium infections 703

A González-Martí nez, MJ Olivares-Duran, Juan Francisco Gutiérrez-Bautista, JA Reguera Márquez, Luis Aliaga, and Javier Rodrí guez-Granger

Leptospira, Borrelia and Treponema 719

Antonio Sampedro and Francisco de Así s Ramirez

Mycoplasma and Ureaplasma 730

Carmen Liébana-Martos

Legionella, Chlamydia and Chlamydophila 737

Elena Cuadros Moronta

Ehrlichia, Anaplasma and Related Bacteria 749

Juan Francisco Gutiérrez Bautista, Francisco de Así s Ramirez, and Anastasia Sampedro Padilla
 General Concepts of Immunity
Nima Rezaeia,b,c, Mona Sadeghalvada,b, and Hamid-Reza Mohammadi-Motlaghd, aDepartment of Immunology, School of Medicine,
Tehran University of Medical Sciences, Tehran, Iran; bNetwork of Immunity in Infection, Malignancy and Autoimmunity (NIIMA), Universal
Scientific Education and Research Network (USERN), Tehran, Iran; cResearch Center for Immunodeficiencies, Children’s Medical Center,
Tehran University of Medical Sciences, Tehran, Iran; dMedical Biology Research Center, Health Technology Institute, Kermanshah University of
Medical Sciences, Kermanshah, Iran
© 2022 Elsevier Inc. All rights reserved.

Introduction to immune system 1

Innate immunity: The first line of defense against pathogens 2
Effector mechanisms of innate immune response 2
Recognition of conserved molecular patterns of pathogens 2
Phagocytosis: A killing mechanism by phagocytes 2
Inflammatory response: A process induced by pathogen recognition and tissue damage 3
Complement system 3
Antibody-dependent cell cytotoxicity (ADCC): An extracellular killing mechanism 4
The components of innate immune system 4
Physical and chemical barriers 4
Innate immune cells 5
Monocyte and macrophage 5
Dendritic cell 6
Neutrophil 6
Eosinophil 7
Basophil 7
Mast cell 7
Innate lymphoid cells (ILCs) 8
Natural killer cell (NK) 8
Natural killer T cell (NKT) 8
Adaptive immunity 8
Adaptive immune cells 10
B lymphocytes 10
T lymphocytes 10
Effector mechanisms of adaptive immune response 10
Antibodies: The effector mechanism of humoral immunity 10
Antigen presentation: A mechanism for T cells activation which is mediated by MHC molecules 11
T cell response: The effector mechanism of cellular immunity 11
References 12

Introduction to immune system

Maintaining homeostasis in response to environmental alterations is a significant challenge for living organisms. Elimination of
invading pathogens and damaged or harmful cellular contents is an inevitable challenge that all organisms have to contend with it.
The body’s defense against the potential dangers and re-establish homeostasis rely on “immune system” which consists of the cells
and molecules responsible for protecting the host body from harmful agents. In this way, the comprehensive and organized
response of immune system components leads to the formation of an “immune response.” Although an ideal immune response
would eradicate the imminent danger and restore homeostasis to affected tissues, its improper activity can also be detrimental and
cause tissue damage. Hence, the regulation of immune responses is crucial, allowing the organism to modulate the intensity and
duration of the responses (Chaplin, 2010).
To provoke an efficient response, the immune system must recognize a wide range of infectious agents named pathogens such as
viruses, bacteria, parasites, and fungi and discriminate them from the host’s own tissues. This process is performed by a variety of
receptors expressed on the immune cells (Medzhitov and Janeway, 1998). Any substances recognized by immune system compo-
nents named “Antigen” and the specific part of an antigen recognized by immune receptors is called “Epitope.” One of the unique
features of the immune system is the ability to distinguish between “self” and “non-self” antigens. To prevent inappropriate
reactions, the immune system must have tolerance against host’s own molecules. This is an important event which states the concept
of “immune tolerance” theory. Disruption of self-tolerance or inappropriate regulation of immune responses could lead to immune
reaction to host cellular components resulting in a state known as “autoimmunity” (Chaplin, 2010).

Encyclopedia of Infection and Immunity https://doi.org/10.1016/B978-0-12-818731-9.00043-4 1

2 General Concepts of Immunity

In overall, immune system can be categorized into two types include “innate” and “adaptive” immunity. The innate immune
system is the first line of defense which quickly activated against pathogens. This type of immune response reacts in the same way to
all foreign antigens, which is why it is named as the “non-specific” immunity. However, an efficient immune response requires the
involvement of adaptive immunity. The adaptive immune system specifically recognizes the antigens and responds slower than
innate immune system. The advantage of adaptive immunity is the production of “memory” cells which can react immediately
against pathogens. Therefore, when a known antigen is encountered for second time, the adaptive immune system can react faster
(McCullough and Summerfield, 2005). In this article, we aimed to describe the basic concepts of immune system including the
important components of innate immunity and adaptive immunity as well as the basic effective mechanisms in order to pathogen
elimination.

Innate immunity: The first line of defense against pathogens

As an old evolutionary defense mechanism, the innate immune system or native immunity, provide the first line of defense against
pathogens. Innate immunity is responsible for respond immediately (within minutes) to microbial components and harmful
agents caused by cellular stress or tissue damage. Innate immune system consists of the cellular and biochemical components that
are present active even before infection. Although some of them including complement system and cellular compartments become
active upon danger exposure. An important feature of the innate immune system is its ability to activate the adaptive immune
system (Tosi, 2005; Beutler, 2004).
Innate immune response provides the immunity against pathogens through several ways including:

1. Pathogens blocking from entering the body.

2. Pathogen uptake by phagocytosis or pinocytosis.
3. Pathogen killing via cytotoxic mediators.
4. Increased recruitment of the other immune cells by production of inflammatory mediators.
5. Pathogen processing and antigen presentation to adaptive immune cells.

Effector mechanisms of innate immune response

Recognition of conserved molecular patterns of pathogens
Innate immune response relies on the recognition of molecular patterns existed on the pathogens or damaged associated molecules
which produced during injury, infection, or disease condition. These molecular patterns are including “pathogen-associated
molecular patterns” (PAMP) such as lipopolysaccharide (LPS) of Gram-negative bacteria, peptidoglycan of bacterial cell wall,
and viral RNA; and “damaged-associated molecular patterns” (DAMP) such as heat shock proteins (HSPs) and high mobility group
box 1 (HMGB1). PAMPs or DAMPs could be recognized by non-specific receptors named “pattern recognition receptors” or PRRs
expressed on immune system components. Several types of PRRs including Toll-like receptors (TLR), nucleotide-binding oligo-
merization domain (NOD)-Leucine Rich Repeats-containing receptors (NLR), retinoic acid-inducible gene 1 (RIG-1)-like receptors
(RLR), and the C-type lectin receptors (CLRs) have been identified which are differently diffused in sub cellular compartments such
as cytosol, cell membrane, and endosomal membrane. Ligand-induced activation of PRRs initiates inflammatory responses
resulting in the promotion of pathogen elimination as well as tissue homeostasis (Mogensen, 2009).

Phagocytosis: A killing mechanism by phagocytes

Phagocytes are the major innate immune cells act as the first line of defense against microorganisms. The important phagocytes are
monocytes and macrophages, neutrophils, eosinophils, basophils, mast cells, and dendritic cells. Anatomically they are abundantly
found in the locations where they encounter most pathogens, such as submucosal tissue of the respiratory system, gut, skin, and
urogenital tract (Rabinovitch, 1995).
The initiation of phagocytosis is dependent on the interaction of microbial surface with certain receptors on the phagocytes.
Subsequently, the bounded microorganism is surrounded by the cell membrane of the phagocytes and internalized in a vesicle
named “phagosome.” Then, lysosomes which are enriched in antimicrobial enzymes are fused into phagosome leading to the
formation of “phagolysosome.” In this context, pathogen killing would occur through the enzymatic process or by the influence of
antimicrobial peptides present in phagolysosomes (Fig. 1) (Flannagan et al., 2012).
Several receptors have been identified that stimulate the phagocytosis and intracellular killing of pathogens. “Dectin-1” is
expressed by macrophages, neutrophils, and dendritic cells which recognizes the polymers of glucose especially on the fungal cell
wall (Herre et al., 2004; Dambuza and Brown, 2015). Furthermore, “Mannose receptor” (MR) is expressed by macrophages and
dendritic cells which recognizes mannose residues in bacteria, virus, and fungi (Ezekowitz et al., 1990). Also, “Scavenger receptor”
(SR) on macrophages identifies different anionic polymers and low-density lipoproteins (LDL) (Peiser et al., 2000).
Phagocytosis could be enhanced by binding of some components named “opsonins” to the pathogen. Immunoglobulins and
complement fragments (e.g., iC3b) are identified as the key opsonin molecules which induce efficient phagocytosis (Rosales and
 General Concepts of Immunity 3

Fig. 1 The effector mechanisms of innate immune system. Ig, Immunoglobulin; ADCC, antibody-dependent cell cytotoxicity; NK, natural killer cell; Eos, eosinophil;
MQ, macrophage; COM, complement.

Uribe-Querol, 2017). Therefore, complement receptors (CRs) and Fc receptors (FCRs) are the other receptors that facilitate
phagocytosis and enhance pathogen killing function (Fig. 1) (Rosales, 2007).
During the pathogen killing by phagocytosis, some toxic compounds are produced that including superoxide ðO2 − Þ, hydroxyl
radical, nitric oxide (NO), hypochlorite (HOCl), and hydrogen peroxide. This process is accompanied by a mechanism known as
the “respiratory burst” which included a transient increase in oxygen consumption by the phagocytes. There is another mechanism
to pathogen take up known as “macro-pinocytosis.” In this way, large amounts of extracellular fluid along with the microorganism
are ingested (Uribe-Querol and Rosales, 2017).

Inflammatory response: A process induced by pathogen recognition and tissue damage

Recognition of pathogens or damage-related molecules elicits an inflammatory response within initial hours of infection that results
in increased infiltration of leukocytes to the affected tissue as well as elevated production of acute phase proteins (APPs). This
response is accompanied by heat, redness, pain, and swelling at the site of infection (Kotas and Medzhitov, 2015; Nathan, 2002).
Interestingly, inflammatory signaling induces blood clotting in small vessels at the site of infection which plays an important role in
preventing the dissemination of infection throughout the body (Levi et al., 2003).
During inflammatory response, immune cells produce a variety of inflammatory mediators including the proteins or glycopro-
teins named “cytokines” and “chemokines.” Interleukin-1 (IL-1), IL-6, and tumor necrosis factor-a (TNF-a) are the most important
pro-inflammatory cytokines produced by innate immune cells. These mediators increase the vascular permeability and the
recruitment of the other innate or adaptive immune cells leading to the promotion of immune response against pathogen or injury
(Gilroy et al., 2004; Headland and Norling, 2015). In this context, inflammatory response induces the expression of cell-adhesion
molecules on the endothelial cells leading to the promotion of leukocytes binding and their migration from blood stream to
affected tissue, a mechanism known as “extravasation.” Meanwhile, Neutrophils are the first leukocytes attracted to the infection site
(Mitroulis et al., 2015).
Surface molecules of microorganisms also activate complement system leading to production of complement-associated
molecules such as C5a or C3a which act as “anaphylatoxins” and increase the recruitment of immune cells to the infection site
(Merle et al., 2015).

Complement system
Complement system as a pivotal effector mechanism of innate immune system is capable of destruction of a variety of microor-
ganisms including bacteria, viruses, and parasites. This system consists of more than 30 soluble and cell surface proteins (Fig. 1)
(Merle et al., 2015). Upon stimulation, these proteins being activated and initiate a cascade which results in:

1. Pathogen destruction by forming pores on the cell membrane of microorganism

2. Increased recruitment of inflammatory immune cells by production of “anaphylatoxins”
3. Enhanced phagocytosis by production of opsonins
4 General Concepts of Immunity

The complement system could be activated by three pathways including:

1. Classical pathway: activated by the immune complexes of antigen-antibody

2. Alternating pathway: activated by the spontaneous hydrolysis of C3 component
3. Lectin pathway: activated by the recognition of certain carbohydrates on the microbial surface

An immune complex which consists of antibodies (IgG1 or IgM) attached to the cell surface of pathogens is capable of activating the
classical pathway. In this manner, C1 complex (C1q, C1r, C1s) binds to the Fc region of antibodies. Upon activation, C1s acts as a
serine protease and cleaves C4 and C2 into C4a, C4b and C2a, C2b, respectively. Subsequently, C4b along with C2a form C4bC2a
complex which interacts with pathogen surface. This complex (C4bC2a) acts as an enzyme named “C3 convertase” which cleaves C3
into C3a and C3b fragments. This step is the point in which all complement pathways converge. C3b fragment covalently binds to
the microbial surface and its association with C4bC2a makes a complex (C4bC2AC3b) named C5 convertase which in turn converts
C5 into C5a and C5b. Binding of C5b to the microbial surface leading to activation of other components including C6, C7, C8, and
C9. Ultimately, polymerization of C9 on the pathogen surface results in the formation of several pores named “membrane attack
complex” or MAC which can directly lyse targeted pathogen. The lectin pathway is similar to classical pathway, but its activation is
antibody-independent. In this pathway, pattern recognition receptors (PRRs), such as mannose-binding lectin (MBL) and ficolins
recognize the carbohydrates on the microbial surface. MBL is complexed with MBL-associated serine proteases (MASPs)-1, -2, and -3
which their activation results in cleavage of C2 and C4, and subsequently the generation of the C3 convertase (C4bC2) of both
classical and lectin pathways. The alternative pathway is activated with the spontaneous hydrolysis of C3 into C3a and C3b. In the
presence of microbial infection, C3b binds to the microbial surface. Subsequently, the association of B and D factors with C3b leads
to additional C3 cleavage and formation of alternative pathway C3 convertase (C3bBb) and C5 convertase (C3bBbC3b). A protein
named “properdin” stabilizes alternative pathway convertases and facilitates alternative activation. Anaphylatoxins including C4a/
C3a/C5a are potent inflammatory molecules which possess several functions such as increases in vascular permeability, leukocyte
infiltration, smooth muscle contraction, phagocytosis and production of inflammatory mediators such as pro-inflammatory
cytokines (Merle et al., 2015).

Antibody-dependent cell cytotoxicity (ADCC): An extracellular killing mechanism

ADCC could be considered as an extracellular killing mechanism resulting in pathogen elimination. This process requires:
(1) immune cells expressing Fc receptor on their surface, and (2) target antigen coated by antibody (Fig. 1). Immunoglobulin
G (IgG) and its receptor named FcgRIII (CD16) play an important role in ADCC activation. NK cells which express high levels of
CD16 are regarded as the key players in this process. However, in addition to NK cells, macrophages, neutrophils, and eosinophils
are able to mediating ADCC (Hubert et al., 2011; Siders et al., 2010; Valerius et al., 1990).
IgG possess a bifunctional structure which is related to the fragment antigen-binding (Fab) and Fc portions of antibody. ADCC is
initiated by the engagement of Fab and Fc regions of antibodies with the pathogen and FcgR on effector cells, respectively.
Subsequently, degranulation of effector cells (mainly NK cell) leading to pathogen lysis (Nigro et al., 2019).

The components of innate immune system

Physical and chemical barriers
The innate immune system includes anatomical and chemical barriers that impede microbes from entering the body. Epithelial cells
which comprise the skin and the lining of the respiratory, gastrointestinal, and genitourinary tracts act as the first line of defense
against pathogens. In addition to forming the tight junctions, epithelial cells produce a large amount of “mucus” enriched
glycoproteins known as “mucins” which traps microorganisms and prevent their dissemination (Fig. 1). Epithelial cells also provide
a chemical barrier by producing various chemical substances with antimicrobial properties. Digestive enzymes and acidic pH of the
stomach, lysozyme, phospholipase A2, defensins, cathelicidins, and histatins are identified as the important chemical barriers to
infection (Schleimer et al., 2007).
“Lysozyme” is a glycosidase enzyme in tears and saliva which has the ability to disruption of peptidoglycan present in the
bacterial cell wall especially in Gram-positive bacteria. Paneth cells, a type of epithelial cells in the small intestine produce several
microbiocidal proteins such as lysozyme and phospholipase A2 responsible to killing the pathogens in the gut lumen (Sukhithasri
et al., 2013).
“Defensins” as the cationic peptides form a pore in the cell membrane of pathogens like bacteria and fungi as well as the viruses’
envelope. Two distinct types of defensins with different activities and structure are implicated in humans including a- and
b-defensins. Neutrophils and paneth cells produce a-defensins. The production of b-defensins is induced by infection considerably
in skin, airway and urogenital tracts (Ganz and Lehrer, 1997; Ouellette and Selsted, 1996).
“Cathelicidins” are antimicrobial peptides produced by macrophages, neutrophils, mast cells, NK, T, B, sebocytes, and epithelial
cells in response to infection (Roby and Di Nardo, 2013).
“Histatins” as the important anti-fungal peptides have a key role in host defense in the oral cavity. Histatins are primarily
produced in response to pathogenic fungi such as Cryptococcus neoformans and Candida albicans (Edgerton and Koshlukova, 2000).
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Exploring the Variety of Random
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 dooty. It nearly breaks
my heart to go so far
away and leave you
behind; but we have to
give up the old farm,
Betsy, we have to give
up the old farm, and I
must find some place to
go to, and something to
do. We must live, Betsy,
—we—must—live,—and I
must find something to
do, to live. I hope to be
able to find work, and
have you to come to
where I am before long.
“I surely can find
something to do some
place. I heerd Jonas
Warner, that rich man in
town, tell a feller the
other day that anybody
could find work that
wanted to work. God
“He drawed me over in his arms and knows, Betsy, I want to
kissed me.” work, and if Mr. Warner
is right, I surely can find
somebody willin to give
me something to do.”
We dident sleep much that nite. Jobe wanted to ketch the five
o’clock train on the C., L. & W. Railroad, and was afraid of
oversleepin hisself. He had to git up airly so as to git to town in time
to ketch it.
That mornin I had his clothes done up in a neat bundle. I had
washed and ironed all his clothes the day before, so he would have
enough to do him till I could go to him.
 He dident eat much breakfast.
He said he “dident feel hungry.”
When he got ready to start he
come up to the winder where I was
a standin, and, seem that I was
choked up, my eyes full of tears,
he drawed me over in his arms and
kissed me; then, turnin, walked out
of the door without sayin a word.
The moon was a shinin bright, and
I stood a lookin at him as far as I
could see him. He was wipin his
eyes and blowin his nose as he
went towards town.
When he was gone from my view
I still stood a lookin for some time,
then sot down and cried, and kept
a cryin every little bit all mornin.
Everything seemed so lonesome
like. Wherever I looked it seemed I
could see poor Jobe a standin there
lookin sad like.
He said he would rite as soon as
he found work. I am lookin for a
letter every day.
Poor Jobe! Little did he think, or
me either, some thirty-six years “He was wipin his eyes and
ago, that in our old age we would blowin his nose as he went
be turned from our home by the towards town.”
law of our country. Little did we
think that when we got old Jobe would have to go hundreds of miles
from home, and out among strangers, a beggin for work to feed us
by.
 “Then sot down and cried, and kept a cryin every little
bit all mornin.”

Jist to think of all the interest money and payments we have give
Congressman Richer—some $3,800 all told. If interest had been less
we would have had our home, and had it nearly paid for, and Jobe
would not be gone out into the world to hunt work. If we had half or
a quarter of that interest money we could buy us a little home to
stay in the few remainin years of our lives.
But, then, interest must be kept up, and the law inforced, so as to
enable Mr. Richer and his likes to live in style and assert the dignity
of their citizenship. It has to be done, no matter if the hardworkin
poor people are turned out of their homes and those that love each
other are parted.
If Jesus was here and a makin laws, I wonder if he would have
interest, and foreclosin, and turnin out, and all that?
 CHAPTER XXXIX.
THE PREACHER AND THE SALOONKEEPER.

M Y heart is so broke that I hardly know how to rite. This is March

3d, and yisterday arternoon they put me out.
I had about give up their comin, and was tryin to feel better, when
all of a suddint I heerd a knock at the door. I opened it, and there
stood three strange men.
Said the one who acted as leader: “Is this where the Gaskinses
live?”
Says I: “One of them is stayin here, and the Lord only knows
where the other one is.”
“I am a deputy sheriff,” says he, “and have orders to set you out.”
Says I: “Where is Mr. Richer?”
“In Washington,” says he.
“Where is his agent—his lawyer?” says I.
“In town,” says he.
“Well, dont they have to be here to put me out?” says I.
“No,” says he; “the law puts you out for them.”
“Well, Mistur,” says I, “couldent you let me stay a little longer?
Jobe’s gone to hunt work and a place to move to. If you will let me
stay, as soon as he finds it Ile go out without your botherin.”
“I cant do it, Mrs. Gaskins,” says he; “the law must be inforced.
The law is no respecter of persons.”
Says I, pleadin like: “You see, I am a old woman, and not stout.
Jobe is away, and I am here alone. If the law is no respecter of
persons, why should it come here and put me out of a home that we
have paid over $3,800 toward, jist to please the man that we have
paid the money to?”
 He shook his head.
“Where are you a goin to put me?” says I.
“I am goin to put you out,” says he; “out in the big road yonder,
off these premises.”
Says I: “Mistur, please dont be so cruel as that. It would kill me to
sleep out there all nite. Please let me stay a little longer—jist a little
longer.”
“No use a talkin,” says he. “Ile have to do as the law says. Its not
me a puttin you out, Mrs. Gaskins—its not me that is cruel. It is the
law, the law, that is doin it.”
“Come on, men,” says he, speakin to the other fellers.
So they come right into the house, the house I had loved so well,
walkin over the floor I have scrubbed on my hands and knees
thousands of times, and begin to tear up my things and carry them
out in the big road.
I jist felt so queer I could hardly breathe.
They tore down my stove and tore up my carpet, and carried out
fust one thing, then another, and sot them down beside the road, till
all I had was out there.
When they got it all out, the deputy come in and says:
“Why dont you go out there where your things are? You have no
right here. You must git out, so I can lock up the house.”
Says I: “Mistur, is Congressman Richer a goin to move in to-nite?”
Says he, sneerin like: “Why, Lord no; Mr. Richer wouldent live in
sich a house as this—he lives in Washington; he lives in a fine
house.”
“Well, then, Mistur, let me stay in here till I hear from Jobe.”
“No,” says he, “you must git out.”
 “They pulled me away from the winder.”

Says I, chokin like: “Mistur, I cant go.”

“Well, youve got to go,” says he. “Are you a goin?”
“I cant,” says I.
“Here, men,” says he, “take her out of here and out yonder, where
she belongs.”
So one of them big men took hold of one arm, and the other hold
of the other arm, and pulled me away from the winder where I was
standin (the same one where I was standin the mornin Jobe left),
and pulled me out of that dear old kitchen door and across the yard
and out into the big road, where they had piled my things, and sot
me down on a chair.
The sheriff had locked the house and follered them out.
When he came out he says, as though he wanted to be friendly:
“Where do you think of goin to, Mrs. Gaskins?”
I looked at him to see if he was crazy or what, but I couldent
speak, I was so full.
Says he: “Do you want the boys to put up your bed for you?”
I nodded my head.
They set my bed up and put two jints of pipe on my stove, and
then got in their buggy and went to town. It was nearly sundown
when they left me.
Soon arter they had gone Tom Osborne come a ridin by and
brought me a letter.
As soon as he said “letter” my heart leapt. I knew it was from
Jobe.
Tom said he was sorry to see me out here in the road, and the
man really shed tears. He lives some eight miles from here, and
wanted me to go home with him for the nite. But I jist couldent go.
So he rode on.
Arter he was gone I got a lamp and sot down by the fire I had
built in the stove, with some quilts around me, to read poor Jobe’s
letter. And every word seemed to be another knife stuck in my heart.
Poor Jobe he is havin it hard too. I jist cried like my heart would
break as I read what he writ. I send it to you to read. I want you to
return it, as it is from the only person in the world that cares for me.
Here it is—you can read it for yourself. You see it was writ at
different times and places.

JOBE’S FIRST LETTER.

Elyria, O., Feb. 22, 1896.
To Betsy Gaskins.
My Dear Wife:—I have put off ritin to you thinkin I would be able
to rite you somethin to make you happy, but to date I cant.
 I got into Lorain the third day arter leavin you. I found a big iron
works there and lots of men at work, but on the sides of the door to
their office and at all the gates around the big fence they have signs
stuck up, readin:
NO HELP WANTED HERE.
I went into their office, and asked them if they couldent give me
something to do.
They said: “No, we have all the men we need.”
I told them how I wanted somethin to do at any price; of our bein
foreclosed and havin to git out and all. They shook their head and
said they “had to turn away hundreds of men every day,” and told
me to “look around,” I “might find work somewhere else.”
So I left and went from one place to another, and everywhere I
went I saw them signs and was told the same thing.
I found lots of men huntin work.
On nearly every street, and down along the river and over by the
lake, were men a campin and a sleepin in railroad cars and
outdoors; cookin by fires built along the banks and on the shore;
“waitin,” they said, “till they could git a job.”
I got my supper with three fellers that nite that done their cookin
that way. They seemed to be nice fellers. They was from different
parts of the country.
That nite I got a bed for fifteen cents, and had forty-three cents
left.
The next day I walked and walked and walked to find work, but
couldent.
At nite I had twenty-four cents left. Not wantin to git clear out of
money, I got into an empty box-car and slept the best I could. It
was cold, and most of the nite I had to walk from one end of the car
to the other, back and forth, to keep myself warm.
So this mornin I come down here to Elyria, and have been from
one end of the town to the other tryin to find work; but nobody
seems to want to hire me.
I find men stayin out around town here too. They say they have
been all over the country, and cant find work anywhere. I dont know
 what I will do. Ile go
over to Berea and see if
I cant find somethin
there. I will not send this
letter till I git there.

“At all the gates around the big fence

they had signs stuck up.”

Cleveland, O., Feb. 26, 1896.

Box-car 1406, Valley Railway.
Betsy:—I am here. I will finish my letter. God only knows what it is
to be out of work, out of money and out of home. I am not well. Ive
had to sleep outdoors, in cars and barns and around lumber piles so
much that I have a bad cold. I have not had anything to eat since
yisterday mornin. This cold weather has nearly used me up. I got
one day’s work cuttin ice, and got a dollar for it. That nite I got me a
warm supper and slept in a bed.
I run out of money at Elyria, and come from there to Berea.
The first beggin I done was from the farmers on the way. I got
one warm meal and a cold lunch. I was in Berea a whole day and
nite without anything to eat, so I jist had to go to beggin agin. I
 went to the Methodist
preacher’s house one of them
real cold mornins. I knocked,
and the preacher come to the
door. I asked him for
somethin to eat. He called to
the hired girl and told her to
hand me a lunch, and went
in, shut the door, and sot
down by the fire. I could see
him a settin there a readin
the Cleveland Leader, with
his feet restin on a plush
foot-stool, and while that girl
was a gittin that lunch and I
was a standin out there in the
wind a lookin at that good big
fire I thought I would freeze.
My teeth shook.
When the girl brought that
“I asked him for something to eat.” lunch I was so cold that I
could hardly take it. It was
two pieces of cold bread, with
some cold beef shaved off and laid between.
I was hungry and tried to eat it; the bites seemed to stick in my
throat, it was so dry and cold. What I did swallow seemed like
chunks of ice in my stomach, and made me colder. I shook from
head to foot. I couldent eat it, I was so cold. So I put what I
couldent eat in my pocket, thinkin I would eat it when I got warmer.
I thought Ide die with cold. No matter how fast I walked, I dident
get warm. I went on and on till I got down where the bizness
houses were. I could smell coffee and warm meat a fryin. It jist
seemed as though I had to go in and take some, but I knew I
darent. It seemed to make me colder. Finally I saw a sign sayin:
FREE HOT SOUP.
 When I got up to it a man opened the door, a sweepin. I stopped,
told him I had no money and was cold, and asked him if I could go
into his place and warm.
“Certainly,” says he, “go right in. Ile be in in a minit.”
I went in—yes, Betsy, went into a saloon, the fust time in my life.
Dont blame me. I had to—I was so cold. The stove was red-hot.
When the feller come in and saw how I was shakin, says he:
“Old man, this is pretty cold weather to be out.”
“Yes,” says I, shiverin.
He brought me a chair and told me to set down. Then he felt my
hands and ears and says:
“Why, you are nearly froze.”
I told him about havin to stay out all nite, and about not havin
anything warm for breakfast, the best I could, I shook so.
He went and got a big woolen cloth, held it to the stove till it got
hot, and wrapped my ears up. Then he went and got a little glass
full of liquor, and told me to drink it and it would warm me up. I told
him I hadent any money, and had never drank a drop of liquor in my
life.
“Well,” says he, “I know you have no money, and, if you had, a old
man like you, in your condition, shouldent pay for it. If you dont
wish to drink it I wont insist, but I thought it would warm you up.”
So he set the glass down on the counter and says:
“Ile make you a hot cup of coffee, and then I think you will feel
better.”
When the saloonkeeper set the glass of whiskey down and went
to gittin me some hot breakfast, I seemed to git colder inside as I
got warmer outside. So, Betsy, I jist made up my mind that Ide drink
that glass of whiskey if it killed me. And I did. Soon after I drank it I
felt a warm feelin inside; and as I sot there it jist seemed as though
I could feel myself a thawin out, with that big fire outside and that
glass of whiskey inside. I sot there till the feller had my coffee and
breakfast ready. It was the best coffee I ever tasted,—though, Betsy,
I always loved the coffee you made,—and the fried eggs and the
ham and the hot cakes jist seemed to melt in my mouth.
 Well, arter I had my breakfast the saloonkeeper came around and
sot down and asked me all about myself, and you too.

“‘Well, old man, sich things hadent ort to

be.’”

And as I told all our trouble, about our foreclosin and sellin out,
and my huntin work and not findin it, big tears would every now and
then leave his big blue eyes and roll down his cheeks, and he kept a
swallerin every little bit. When I had told him all, says he:
 “Well, old man, sich things hadent ort to be.”
So, when I got ready to go, he shook my hand and wished me
good luck in findin work; and when he took hold of my hand I felt
somethin hard in his, and when he let go I had a silver dollar in
mine. I handed it back to him, and told him I dident know as I could
ever return it to him.
“No matter, pap,” says he, “keep it. If you are never able to return
it, all right, and if you are able and never see me, ‘do unto some
other human brother as I have done unto you,’ and the debt will be
paid. Times are hard, and I have sich high taxes to pay that it makes
money scarce with me, or I would give you more. I hate to see you
go out in this cold; you are welcome to stay if you wish.”
But, Betsy, I was so anxious to find work and git a place for you
that I couldent stay. So that day and nite I made it to here. This is a
big town, but so far I have found no work.
Your lovin husband,
Jobe Gaskins.

When I got done readin that letter I was cryin out loud. Poor Jobe.
I wonder where he was last nite.
Oh, how I love that man that took Jobe in and warmed him and
fed him!
I love him though he is a saloonkeeper. I could throw my arms
around his neck and cry on his shoulder with love for him and for his
kindness toward Jobe.
Well, this mornin the world seems strange to me. Last nite arter I
had gone to bed and could look up in the clear sky at the bright
stars, it jist seemed to me, while I laid there in my bed beside the
big road, that every star was a eye lookin down on me with pity.
And, thinkin that they looked that way, I was not a bit afraid and
went to sleep, and slept till daylite.
Hopin God will forgive them for makin and havin laws to put sich
people as me out of home, I am
Your troubled and homeless
Betsy Gaskins.
 CHAPTER XL.
“THEM ROOMS.” THE “DIRECTOR OF CHARITIES.”

T HAT mornin arter I wrote you the last time—arter I had built me a
fire in my stove and got my breakfast and washed up my dishes
and made my bed—I sot down on a chair out there by the big road.
I never felt so queer in all my life. Not a sound could be heard,
except over on the hill near Jake Stiffler’s I could heer a cow a
bawlin. It was awful lonesome. No one to speak to, nothin to look
at, except my things piled up there beside the road.
I couldent help thinkin of poor Jobe—his beggin, and bein cold,
and starvin, and sleepin in box-cars, and sich.
Well, arter I had sot there a while a thinkin, I felt so bad that I jist
thought I would go up to the house and take a look at them rooms
and the place we had so long loved as our home.
I felt afraid like to go, but I thought it might cheer me up to look
into them rooms that I had cleaned and papered and swept—the
rooms where Jobe and me had set in and slept; the rooms that had
sheltered us in sickness and in health.
So I jist throwed a shawl over my head, and walked up the walk
that I had walked up thousands of times.
There were the currant bushes, the lilac, the dead poppy stalks.
And all the weeds and posies, that used to appear to wear a smile
for me, now seemed to turn from me as if to say, “We haint yours
any more. You have no bizness here now.”
And as I looked at them and felt that feelin, a lump would raise up
in my throat, no matter how much I swallered and tried to keep it
back.
 Well, I walked on until I
got up to the kitchen winder.
When I got there it jist
seemed that I couldent look
in, but, knowin I had come
there to see them rooms,
half afraid like but
determined, I slipped over
and put my face agin the
glass.
Everything was silent and
still. There was my kitchen,
all empty. Not a thing to be
seen but that dear old
kitchen—empty—no stove,
no table, no chairs, no
nothin. There was the
winder where I stood cryin
the mornin Jobe left. There
by that winder I had set a
combin my little Jane’s hair
years ago, while she drew
pictures on them same
winderpanes with her little
fingers. There were the nails
Jobe had drove in the wall
when we fust moved in;
there was the same floor
over which we had walked
for years. Oh, how I longed “I slipped over and put my face agin
to be a walkin over it agin! I the glass.”
was locked out—I couldent
git in.
So I went from one winder to another, lookin in at them rooms.
There was the same grate that had warmed us; there in that corner,
evenin arter evenin, Jobe had set and studied; there in the other
corner I had set and knit, or set and read. It seemed that I could
see Jobe there now. Oh! how I would love to see him there. Poor
Jobe! I wonder if he thinks of the evenins weve spent beside that
fire together. There was our bed-room—empty, silent and still—no
bed, no nothin. There in that room I had set, nite arter nite, with
little Jane when she was sick; there she had throwed her little arms
around my neck and put her fevered face agin mine the last time.
From that room Ellen Jane Moore had carried her arter she was
gone. It was empty now. I was locked out. I couldent go in.
Turnin from them rooms, I walked around the yard, lookin at the
fence, the well, the coal-house, and the things that had been mine.
Then, comin to the front yard, I come to the little white rose-bush; it
seemed to look at me pleadin like. I started to go on, but I couldent.
That rose-bush seemed to call me back. So I jist got me a sharp
stick and dug it up, and took it down to where my things were and
wrapped it up in a cloth.
When I got back to the big road, and was settin there wonderin
what Ide do, how long Ide have to live there in the big road, where
Ide go to and sich, Constable Bill Adams come a ridin by.
When he got up to me, says he:
“Why, Mrs. Gaskins, what are you a doin with all this stuff piled in
the road?”
“Ime livin here,” says I.
“Well, youle have to git this stuff out of the road,” says he. “You
darent obstruct the public highway. Its dangerous to have a pile of
stuff like this in the big road; its liable to scare horses, and
somebody might git hurt or killed. Its aginst the law, Mrs. Gaskins,
its aginst the law, and you will have to move it.”
“The law put it here,” says I.
“No matter,” says he; “youle have to git out of here, or youle be
arrested.”
“Where will I put it?”
“How do I know?” says he. “Youle have to look out for that
yourself. Git it out of here, and that mighty quick, or you will git
yourself into trouble.”
And he rode on towards town.
 Well, as he rode away I sot down and begin to think. Here I was,
a old woman, set out in the big road by the Law—put out of the
house we had paid $3,800 towards; the house empty, and now
comes the Law and orders me to even git away from where the Law
had put me. What to do I dident know. I jist sot there a cryin and
helpless, when I heerd wagons comin down the road. I looked up,
and there come two wagons and four men down the hill.
They drove up and stopped, and there was Tom Osborne, and
Charley McGlinchey, and that fat black-smith, and Jones the baker,
all from Mineral Pint. They had come to move me.
Tom Osborne had went home the night before and told them
about me bein put out in the big road, and they went together and
got teams and come and moved me to town here.
They seemed to be nice, kind men, but talked like them Populists.
They dident talk much to me, but I heerd them talkin to each
other, sayin: “Its a shame,” “a disgrace to civilization,” “wrong,”
“wouldent be if the people could borrow money from the
government like they do in Switzerland,” and all sich. They even
said: “The time haint fur off when it can be done, and the likes of
this wont be.” And then they said a good deal agin the money power
and polerticians, and sich, until I was glad Jobe wasent there to flare
up. I was glad he wasent there, though Ide give the world to know
where he is, or to have him with me.
Well, they brought me to town and rented me this house here at
1412 West Front Street, and paid the rent for a month; then two of
them drove off, and soon brought me a load of coal. While them two
were gone for the coal the other two set up my stove, and fixed up
my bed, and set things around in pretty good shape for men; then,
wishin me good luck, and hopin Jobe would soon git work and I
would git to go to him, they drove off. They all looked pityin like as
they left.
I went to the post-office the next mornin to tell them I had
changed my place of livin. I got this letter from Jobe. It jist seems
there is no end of trouble for the people who are poor.
Poor Jobe, how my heart bleeds for him. Here is his letter. Read it
for yourself:
 JOBE’S SECOND LETTER.

Cleveland Work-house,
Cleveland, O., March 5, 1896.
To Betsy Gaskins.
My Dear Wife and Only Friend:—I am here in this prison—put here
by the law. God only knows my feelins. I am not a criminal. Ive done
no wrong. Betsy, don’t blame me. Pity me. I am a old man. I have
worked hard. Ive been honest. Ive tried to do right. To-day I am in
prison, wearin stripes. I was hungry. I had no money. I asked for
bread. They arrested me.
It was day before
yisterday. I had
hunted for work all
day. I had had nothin
to eat for a whole
day and nite. I was
passin up Ontario
Street, near Hull &
Dutton’s big clothin
store. I saw a well-
dressed man, with a
high silk hat on, with
a hand full of paper
money, talkin loud
and offerin to bet
$500 that McKinley
would git the
delegates from
Allegheny County.
There were several
fellers standin there a
listenin and talkin,
and two policemen. I
stepped up and asked the feller with the money if he could give me
enough to git me a supper and bed. I was so hungry and nearly sick
by sleepin outdoors.
The feller turned around and looked black at me. Then, turnin to
the policemen, he ordered them to arrest me, sayin:
“Ime d—d if I dont intend to break up this beggin on the streets.”
The policemen took hold of me and jerked me out of the crowd
and pulled me down Champlain Street hill to the city prison, and
locked me in a iron cage.
I asked one of them who the big man was that ordered me
arrested. He said it was “the Director of Charities, one of the leadin
city officers.”
You may have read in the papers of him a havin a tramp arrested
for askin him for somethin to buy bread with.
That tramp, Betsy, was me.
They say he gits $5,000 a year for bein “Director of Charities.”
Well, they tried me next mornin and found me guilty.
I am up for ten days. I cant find any work or a place for you till I
git out.
They brought me out here in a wagon with a cage on it. They call
it the “Black Mariar.” There was a lot of us in it. Betsy, pity me. Dont
blame me.
Your lovin husband, Jobe Gaskins.

Mistur Editure, I cant comment. I feel so bad.

 CHAPTER XLI.
A SORE HAND.

I AM sick. I have been sick since day before yisterday. I have a high
fever. My head bothers me. I cant rite. Here is another letter I got
from poor Jobe. Oh! how I wish he was here. I know he would care
for me and watch over me and do for me while Ime sick. Read his
letter and return it. They seem so near to me. I havent been able to
be out of bed much to-day. If Jobe was only out of that dreadful
place.

JOBE’S THIRD LETTER.

Cleveland Work-house,
Cleveland, O., March 9, 1896.
To Betsy Gaskins.
Dear Wife:—I got your letter yisterday. I cant tell you how I felt
when I read of them a puttin you out.
Betsy, I little thought, the day you stood beside me and become
my wife, that the time would come when you would have to sleep
outdoors in the big road.
I felt then, Betsy, as though I was strong enough, and God knows
I was willin, to provide a home for you as long as we both lived.
Dont blame me, Betsy. Ive done the best I could. You know Ive
worked hard, and we have lived savin, but by some unknown reason
all I have aimed is gone. Mr. Richer has $3,800 of it. Ive done the
best I could.
I have to work hard here in this place, but Ime not complainin,
nor wouldent complain if I was gittin paid for what work I do, so
that I could help you.

“I have to work hard in this place.”

Ime a wheelin coal to the furnace and a wheelin hot cinders away.
It keeps me bizzy.
There are lots of men in here. A great many for beggin—jist as I
am. Betsy, dont let the neighbors know they have me locked up. I
feel so disgraced.
I feel that if that “Director of Charities,” that had me arrested and
put in here, had known that I had feelins; if he had known that I
was a honest old man; if he had thought of the difference between a
old man, hungry, away from home and out of money—I say, Betsy, if
he had thought of the difference between sich a man as I was and a
man drawin $5,000 a year as a leadin city officer, like hisself, I dont
think he could have had the heart to have had me arrested and sent
to prison.
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