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IMMUNOLOGY
ABOUT THE AUTHOR
Dr S R Ramesh is currently serving as Professor in University of

Mysore, Mysuru, Karnataka. After obtaining PhD degree from
University of Mysore in 1980, he briefly served as Research
Officer at the ICMR institution Vector Control Research Centre,
Puducherry. His interest and passion for teaching prompted
him to come back to the Department of Studies in Zoology
at Manasagangotri, University of Mysore, Mysuru, as a Lecturer
in 1981.
Dr Ramesh has also worked as post-doctoral fellow at Institut
für Genetik, Ruhr Universität Bochum, Bochum, Germany
with fellowship from Deutscher Akademischer Austauschdienst
(DAAD), Germany. Later, he worked as visiting fellow at Institut
für Genetik, Ruhr Universität, Bochum and at Institut für Genetik,
Heinrich-Heine Universität, Dusseldorf, Germany. In recognition of
his post-doctoral work, the Deutsche Gesellschaft für Technische Zusammenarbeit (GTZ), Germany
supported him to continue his research work that was initiated in Germany by providing laboratory
equipment when he came back to India.
With research grants from University Grants Commission, Department of Science & Technology
and Department of Atomic Energy, he chose to work on Drosophila as a model organism to study
biochemical genetics of larval and pupal salivary gland proteins, adult male accessory gland secretory
proteins, mutations and behavioral differentiation. Five years ago, he initiated work related to learning
and memory in Drosophila and also on screening and understanding the influence of phytochemicals
on biochemical and genetic system of normal, transgenic Alzheimer and Parkinson Drosophila models.
He has authored 85 publications including research papers and research reports in reputed national and
international journals, in addition to many popular science articles in Kannada.
He has served as a chairman/member of many committees of the University Grants Commission
(UGC), New Delhi. He has also served as a member of Academic Council, Faculty of Science &
Technology, Institutional Human Ethics committee, various committees of University of Mysore and
as a member of Institutional Bio-safety Committee of CFTRI, Mysuru.
Dr Ramesh holds vast postgraduate teaching experience of over 34 years and has offered courses
in Immunology, Cell Biology, Cytogenetics, Genetics, Molecular Biology, Animal Physiology,
Environmental Biology and Animal Behavior. He has delivered many special lectures in different fields
of modern biology at different universities and teacher training programs in various institutions.
IMMUNOLOGY
S R Ramesh
Former Professor
Department of Studies in Zoology
University of Mysore, Mysuru, Karnataka
McGraw Hill Education (India) Private Limited

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Immunology
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Dedicated to
My Alma Mater
University of Mysore
on the occasion of its Centenary Year
&
My Family
for their support and encouragement
&
Faculty and Young Students
who wish to become familiar with immunology
CONTENTS
About the Author ii

Foreword xiii
Preface xv
Prologue… 1
1.1 Introduction 9
1.2 Types of Immunity 10
1.3 Innate Immunity 11
1.4 Specific Acquired Immunity (Adaptive Immunity) 21
1.5 Interaction Between Innate and Adaptive Immunity 25
Summary 27
Short Answer Questions 28
Essay Type Questions 29
2.1 Introduction 30
2.2 Lymphatic System 30
2.3 Primary Lymphoid Organs 32
2.4 Secondary Lymphoid Organs 34
2.5 Germinal Centres 38
2.6 Tertiary Lymphoid Organs 39
Summary 40
3.1 Introduction 42
3.2 Hematopoiesis 43
3.3 Phagocytes 45
3.4 Life History of Macrophages 46
3.5 Functions of Monocytes and Macrophages 47
3.6 Lymphocytes 49
viii Contents
3.7 Dendritic Cells 55

3.8 Basophils 58
3.9 Eosinophils 59
3.10 Study of Human Leukocytes 60
Summary 61
4.1 Introduction 63
4.2 Classification of Antigens 68
4.3 Haptens 68
4.4 Superantigens 69
Summary 70
5.1 Introduction 72
5.2 Structure of Immunoglobulins 73
5.3 Antigen-Antibody Interactions 81
5.4 Antibody Dependent Cell Mediated Cytotoxicity 82
Summary 84
6.1 Introduction 86
6.2 Development of B Cells 89
6.3 Genetic Basis of Immunoglobulin Diversity 93
6.4 Clonal Selection and Cellular Production of Antibodies 105
Summary 108
7.1 Introduction 110

7.2 T Cell Development and Maturation 111
7.3 Structure of TCR 114
7.4 Organization of TCR Genes and Their Rearrangement 115
7.5 T Cell Selection 119
7.6 Effector T cells 121
7.7 Other T Cell Types 127
7.8 TCR and MHC Restriction 129
Summary 131
Essay type questions 132
Contents ix

8.2 MHC Molecules 133
8.3 MHC Polymorphism 140
8.4 Antigen Recognition 140
8.5 Antigen Processing and Presentation 146
8.5 Cross-presentation of Exogenous Antigen 153
8.6 Immunological Synapse 153
8.7 Recognition of Endogenous and Exogenous Antigens 154
8.8 Lipid Antigens 155
Summary 156

9.2 Functions of Complement System 166
9.3 Disorders of Complement System 167
9.4 Classical Pathway Deficiencies 168
9.5 Deficiency of Control Proteins 169
9.6 Complement Receptors 172
9.7 Anaphylotoxin Receptors 172
9.8 Receptors of C3b and its Derivatives 174
Summary 176

10.2 Properties 178
10.3 Types of Cytokines 184
10.4 Cytokine Receptors 189
10.5 Cytokines and Inflammation 192
10.6 Cytokine Therapy 195
Summary 196

11.2 Immunological Tolerance 199
11.3 Autoimmunity 202
11.4 Classification of Autoimmune Diseases 204
11.5 Infectious Agents v/s Autoimmunity 214
x Contents
Summary 216

12.2 Primary Immunodeficiencies 218
12.3 Secondary Immunodeficiencies 231
12.4 Acquired Immunodeficiency Syndrome (AIDS) 232
Summary 242

13.2 Type I Hypersensitivity 246
13.3 Type II Hypersensitivity 257
13.4 Type III Hypersensitivity 261
13.5 Type IV Hypersensitivity 263
Summary 267

14.2 Vaccination 271
14.3 Types of Vaccines 274
14.4 How Do We Get Immunized After Vaccination? 290
Summary 292

15.2 Polyclonal v/s Monoclonal Antibodies 294
15.3 Hybridoma 296
15.4 Applications of Monoclonal Antibodies 300
15.6 Advantages and Disadvantages of Monoclonal Antibodies 307
Summary 308

16.2 Occurrence of Cancer 310
16.3 Tumor Antigens 315
16.4 Antitumor Immune Response 318
Contents xi
16.5 The Paradox: Tumors Elude the Immune System 318

16.6 Cancer Immunotherapy 319
Summary 322
Types of Grafts 325
Radioimmunoassay (RIA) 330

Enzyme Linked Immunosorbent Assay (ELISA) 330
Immunoblotting 333
FOREWORD
I was delighted when Dr Ramesh asked me to write the foreword to his book. Dr Ramesh and I were
postgraduate students in the late 70s in Manasagangotri Campus, University of Mysore, Mysuru, Karnataka
which is a wonderful setting for higher learning. Although we came from different disciplines and
backgrounds we always had interesting, intellectual interactions and that continue now also.
Immunity is the cornerstone of existence of all organisms including humans. In nature, every individual
organism or their community is surrounded by pathogenic and non-pathogenic organisms. We all interact,
defend and/or collaborate with a plethora of micro- or macro-organisms in our environment. It may be
alarming to note that despite our obsession to cleanliness and personal hygiene, every human individual
has more non-human DNA/cells than human DNA/cells. Thus, we are constantly exposed to non-self
environment; our health and disease conditions are dependent on continuing ‘conversations’, ranging from
simple interaction to active war. To participate in these ‘conversations’ all higher organisms including
humans have evolved various strategies of innate and adaptive immunities. The science of understanding
these crucial armaments and strategies is called ‘Immunology’.
Dr Ramesh has been teaching immunology to postgraduate students at the University of Mysore, for
more than two decades. His teachings always made significant impact on young minds, particularly in
this extremely interesting system of complex cellular and molecular network as well as protein-protein
interactions involved in molecular recognition discriminating self from non-self. He has put in lot of efforts
in simplifying and explaining various concepts, phenomena and strategies for easier comprehension. This
book showcases his teaching talent and highlights his passion to train the next generation and get them well-
educated in this interdisciplinary field.
The book covers (a) both fundamental and cutting-edge information in innate and adaptive immunity;
(b) all cellular and protein components and their complementary and collaborative interactions;
(c) their role and implications in autoimmunity, immunodeficiencies and hypersensitivity; and (d) their
applications as in monoclonal antibodies and tumor biology. The key features of the book include Learning
Objectives at the beginning of each chapter, ample illustrations for easy comprehension of the concepts and
mechanisms, boxes highlighting key facts and details, summary and, my favourite, several short answer
questions and essay type questions for students that cover various aspects described in the chapter. The
multiple choice questions at the end of the book, as the icing on the cake, will enhance and test their
understanding. Finally, there are two important appendices; one on transplantation and immune suppression,
and the other on immunoassays.
Overall, I believe this will be an excellent text book for undergraduate and postgraduate students and also
act as a reference book for students of medical and paramedical courses as well as for scientists entering the
field of immunology.
DR R MANJUNATHA KINI
Professor
Department of Biological Sciences
National University of Singapore
PREFACE
Immunology, the study of immune systems, originated when Edward Jenner demonstrated that the
incidence of smallpox can be prevented by introducing a small dose of fluid from cowpox pustule
into people. A greater push for immunology to develop into a distinct branch of biology came from
the work of Louis Pasteur who showed that not only smallpox, but also the incidence of variety of
diseases both in animals and humans can be prevented by immunization with vaccines. Since then,
immunization has saved mankind from sufferings and death emanating from epidemic and debilitating
diseases. Immunology is a field of biology and knowledge of this branch of study has grown leaps and
bounds due to development and advances in biochemical and molecular techniques. Incredible insights
into the nature and molecular mechanisms of interactions that occur between tissues and cells, within
the cells as well as between the cells and, molecules of the immune system have been the basis for rapid
advances of not only immunology but also medicine and pharmacology.
In the beginning of an immunology course, my students often used to express that immunology is
tough and complicated but as they become familiar with a reasonably good comprehension of the basic
concepts, they have always found that “Immunology is Amazing”. These interactions prompted me to
transform my lectures into a book. The content and language in this book has been kept simple for easy
comprehension of concepts, processes, mechanisms and applications that beginners always need. This
book will also be helpful for teachers who wish to become familiar with immunology for graduate or
post-graduate classes and also for venturing into higher levels of learning in immunology.
This book has been organized into two parts. The first part “Immunobiology” comprises 10 chapters
focusing on different components of the immune system. A clear comprehension of the content in
this part will create a mindset to understand and think further about the contents of second part,
“Immunology and Human Health”. The second part comprises 6 chapters that deal with importance
and relevance of the immune system in human health. The last chapter of second part is an overview
on immune system and cancer.
Every chapter begins with an introductory paragraph. Subsequently in addition to basic information,
advanced information distilled from recent review articles has been included with an intention to trigger
thinking in the young minds and to enable them to realize its research and application potential. There
are 78 box items altogether. In every chapter the box items provide clarity of a concept or additional
information in the same field or related field depending on the context. The content in this book has
been further supplemented with tables and figures for providing clarity. Additionally, two appendix
items are included to make the book more comprehensive.
The important pedagogical features are as follows:
● Case studies given in boxed items
● Important terminologies highlighted
● Illustrations: 240
xvi Preface
●Chapter end questions: 326

●Objective type questions: 160
My sincere thanks are due to all the scientists and organizations who have given permission for using
their original figures, which have enhanced the scientific content in this book. I am greatly indebted to
my family members who always were the source of my inspiration. I am thankful to my colleagues,
well wishers and friends, in particular Dr M S Basavaraj, Chief Medical Officer, University Health
Centre and Dr T Shivanandappa, CSIR Emeritus Scientist who always encouraged me by discussions
over the issues related to health and contemporary biology. I acknowledge the assistance of Dr M K
Ramakrishna and Ms Sadhana Mutalik during proof reading of the manuscript.
The support of University of Mysore in my academic endeavors is sincerely acknowledged.
I am extremely happy to contribute this book to the academia at a time when the University of
Mysore is celebrating its centenary year.
Special thanks to the reviewers mentioned here for their valuable feedback and suggestions.
J Kamaleswari
Pacchyappa’s college, Chennai, Tamil Nadu
RC Rajkhowa
Cotton College State University, Guwahati, Assam
NK Ganguly
National Institute of Immunology, New Delhi
RP Singh
Indian Institute of Technology, Roorkee, Uttarakhand
It is my pleasure to work with the McGraw Hill Education team headed by Ms Vibha Mahajan and team
members–Vaishali Thapliyal, Suhaib Ali and Kritika Lakhera, who readily responded to all my queries
that enabled me to finalize this book.
S R RAMESH
Publisher’s Note
McGraw Hill Education (India) invites suggestions and comments from you, all of which can be
sent to info.india@mheducation.com (kindly mention the title and author name in the subject line).
Piracy-related issues may also be reported.
PROLOGUE…
Edward Jenner’s pioneering contributions enabled the eradication

of smallpox from this earth. It is regarded that Jenner’s work laid the
foundation for the establishment of Immunology, its subsequent
development, and progress. Hence, Jenner is also regarded as the
father of Immunology. Smallpox over many centuries devastated
mankind. In the 18th century in Europe, around 4,00,000 people
died of smallpox annually. Those who could recover and survive
were badly disfigured, which was much more frustrating and
agonizing throughout their life. Further, one-third of the survived
lost their vision. Hence, smallpox was otherwise also called
“speckled monster” in 18th century England.
Edward Jenner, a doctor by profession in a county namely
Gloucertershire, in England was also interested in varied branches
of science. He studied geology, conducted investigations on
behaviour of cuckoo, and also carried out experiments on human Edward Jenner (1749-1823)
blood. When he was 13 years old, he was apprenticed to a surgeon (foundersofscience.net)
and apothecary in a place near Bristol. He was fascinated by the
belief that milkmaids in some way remained protected from dreadful smallpox. A remark that he
overheard a milkmaid saying, “I shall never have smallpox for I have had cowpox. I shall never have an
ugly pockmarked face” made him to think loudly and believe that there was some connection between
the milkmaids getting the mild cowpox infection and their resistance to life threatening smallpox.
What is cowpox?
Cowpox is a viral skin disease in cows caused by cowpox virus. The affected cows develop blisters
on udders. The virus is transmitted from cows to humans when the udders are touched while milking
the cow. The symptoms of cowpox infection include appearance of red blisters on the hands of
infected person.
Jenner developed a theory that cowpox not only protected the people against smallpox but also could
be transmitted from one to another as a deliberate mechanism for protecting an individual from smallpox.
Jenner decided to try out the theory he had developed. Jenner met a young milkmaid Sarah Nelms, who
had fresh cowpox lesions on her hands and arms. He persuaded her and collected matter from cowpox
blisters and inoculated it into a young boy James Phipps on 14 May 1796; who subsequently developed
2 Immunology
Victims of smallpox: Left: Extent of suffering during the disease (Content provider: Centers for
Disease Control and Prevention), Right: disfigurement after recovery (Content provider: Centers for
Disease Control and Prevention/Dr. Robinson).
mild fever and discomfort in the axillae. Jenner repeated this process a number of days with a gradual
increase in the amount of matter he introduced into the boy. On 1 July 1796 Jenner inoculated the boy
again the matter and this time not with the matter from cowpox lesion but with the matter from a fresh
smallpox lesion. Phipps never caught smallpox and
Jenner concluded that protection was complete. Thus,
Jenner established a method to prevent the incidence
of disfiguring and often fatal disease smallpox and it
took him twenty years of observation, analysis, and
experimentation by the virtue of which, the current
practice of vaccination took its origin. Jenner decided
to call this new procedure as vaccination (Latin: vacca
= cow) in honour of the part played by Sarah and her
cow ‘Blossom’ in his research. In 1798, after adding
few more cases to his initial experiment, Jenner
published his work as a small booklet titled “An Inquiry
into the Causes and Effects of the Variolae Vaccinae,
a disease discovered in some of the western counties
of England, particularly Gloucestershire and Known Edward Jenner introducing fluid from a
by the Name of Cow Pox”. Jenner’s discovery was cowpox blister into the arm of James Phipps
so successful, that in 1840 the government that was (sblazak.wordpress.com)
Prologue… 3
ruling at that time banned any other treatment for smallpox other than the one described and established
by Jenner. Jenner’s method of preventing the incidence of smallpox spread quickly throughout Europe.
It was nearly 100 years, before the procedure of
vaccination was applied to other diseases also. The French
scientist Louis Pasteur was responsible for initiating such a
work by taking the lead from Edward Jenner. Louis Pasteur
was born in Dôle, France. At the age of 20, he received his
bachelor’s degree in science. At Écôle Normale, Paris he
worked on crystallography for his doctoral degree. When he
was 29 years old, he became the chairman of the department
of chemistry at the University of Strasbourg, Strasbourg,
France where he began studying fermentation. At the age
of 32, Pasteur became professor of chemistry and dean of
sciences at the new University of Lille. At this time, Lille
was the centre of alcohol manufacture in France. Soon after
his arrival at Lille, a producer of vinegar requested Pasteur
for help. The vinegar producer could not understand why his
vinegar made from beet juice sometimes spoiled and wanted
to know how to overcome this problem. Pasteur conducted
a microscopic examination of the beet juice and found that Louis Pasteur (1822–1895)
it contained alcohol and yeast. He suggested that the yeast (www.gardenofpraise.com)
was causing the beet juice to ferment. He also demonstrated
that killing of the yeast without spoiling the product is possible by controlled heating of the beet juice.
This process, called “pasteurization,” was widely employed to preserve a number of foods such as
cheese and milk. Pasteur’s discovery brought tremendous improvements in fermentation and heralded
a new era in the brewing and wine industries. Pasteur was an accomplished scientist. In 1857, he was
appointed Director of Scientific Studies at the Ecôle Normale in Paris.
The extraordinary contributions of Louis Pasteur were development of the germ theory of disease
and the use of vaccines to prevent the diseases. Experiments in his lab on vaccination as a deliberate
attempt to prevent the incidence of a disease began in 1881. At that time, chicken cholera (caused by
what we call today Pasteurella multocida) was a serious problem for farmers. The rapid spread of this
disease would have wiped out the entire flock in just three days. Pasteur had identified the cholera
bacillus and was growing it in pure culture. When injected with the culture, the chicken invariably died
in 48 hours. Unexpected discoveries by accident are not new to science. An incident that occurred in the
laboratory of Louis Pasteur led to the discovery of a vaccine for chicken cholera. Charles Chamberland,
who was working with Pasteur, forgot to inject the ‘disease’ into some chickens and went away on
holiday. When he came back, he saw the jar of bacteria placed on a side in the lab and realized that
he did not do the job that was told. He thought he would inject it into the chickens anyway and made
injections. He was amazed to observe that none of those chickens died; which usually would have
died in 48 hours. He reported his observations to Pasteur. Chamberland was asked to repeat what he
had done but with a fresh culture of chicken cholera germs. In the next set of experiments, they had
two groups; the first group consisted of the chickens that were already injected with old culture and
another group that were not (new batch of chickens). Chamberland injected fresh cultures into these
two groups of chickens. The result was again amazing; the chickens that were previously injected with
the old culture survived; while the other group of chickens that were previously never injected with old
cultures died. So to say, the chickens that were inoculated with the old culture had become resistant
4 Immunology
(immune) to chicken cholera. Pasteur was of the opinion that their bodies had used the weaker strain
of germ to form a defence against the more powerful germs in the fresher culture. When Pasteur saw
these results he realized that he was repeating the work of Jenner that was done 85 years earlier; which
had enabled Jenner to confer immunity to smallpox in humans by vaccinating individuals with a mild
form of cowpox. Pasteur then prepared nonvirulent (attenuated) cultures of chicken cholera vaccines by
growing the cholera bacillus at 42°– 43°C; at which temperature the bacillus is rendered non-infectious.
With this, they had also discovered that a weakened form of a disease can act as a vaccine.
Pasteur further thought, if attenuated cholera bacillus could render chickens resistant to the disease,
injection of an attenuated anthrax bacillus must render the sheep immune to anthrax. By various
techniques involving oxidation and aging, Pasteur found that anthrax vaccines indeed prevented anthrax
in laboratory trials. In April 1881, Pasteur announced that his team was successful in founding a way to
weaken anthrax germs and so could produce a vaccine against anthrax. Overwhelmed by the successes
with anthrax and fowl cholera diseases, Pasteur over the next 2–3 years got involved in identification
and isolation of microbes for many other diseases; including swine erysipelas, childbirth fever, and
pneumonia. The most famous success of Pasteur’s research was the development of a vaccine against
rabies. From the experimental observations, Pasteur conceived the idea and provided evidence that
pathogens could be attenuated or weakened or disabled from causing the disease by exposing them to
environmental insults such as high temperature, oxygen, and chemicals. Though Pasteur proved that
vaccination provided immunity, he did not explain how it worked; may be Pasteur’s priority was to
develop vaccines for a variety of diseases, not to look into the mechanisms of how it worked. At the
age of 46, Pasteur suffered a serious paralytic stroke. He died in 1895 after suffering additional strokes.
He was buried, a national hero, by the French Government. Thus, Edward Jenner laid the foundation
for immunology and Louis Pasteur was the first to initiate the development of the field of immunology,
which now has grown boundless to encompass a variety of disciplines that are related to human health
and welfare.
A group of scientists including Paul Ehrlich, Elie Metchnikoff, Jules Bordet, Emil von Behring led
by Louis Pasteur were responsible to presume the presence of a system in the body which was believed
to defend or protect the body from the invasion and attack of pathogens. They called it as immune
system; which literally means the system that exempts i.e. the system that exempts the body from
the disease. In other words, the immune system was thought to be responsible for providing absolute
protection against an injurious or a disease causing agent.
The immune system is usually compared to a fort and its various cells to soldiers. Now, we know
that it is a remarkable defense system that has evolved in vertebrates to protect them from invading
pathogenic microorganisms. The system is scattered all over the body from top to toe comprising
various organs, tissues, cells, and molecules. The system is capable of recognizing limitless variety of
foreign invaders and has evolved mechanisms to get rid of them. The efficiency of an immune system
lies in its ability to distinguish the “self” and “nonself” molecules. A variety of cells and molecules
precisely orchestrate the complex events that are associated with identification, neutralization and/or
destruction of the invader.
Immunity is the state of protection from infectious disease. It has both innate and adaptive
components. The innate immunity refers to the inherent potential of the system to impart resistance to
pathogens that invade an individual. This type of immunity is accomplished by anatomic, physiologic,
endocytic and phagocytic as well as inflammatory barriers to name a few. The adaptive immunity is an
acquired immunity. It is specific and reflects the presence of a functional immune system, capable of
specifically recognizing and selectively eliminating foreign bodies including microbes. Unlike innate
immunity, acquired immunity is characterized by specificity, diversity, self/nonself recognition, and
memory.
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interesting instances, and we shall deal with them presently. But
before we proceed to discuss them let us turn back for a moment to
Robert Fulton. After he had at length established the steamboat as a
thoroughly sound concern in America we find him not unnaturally
sighing for other countries to conquer. Accordingly he set his mind
on introducing the steamboat not merely on the chief rivers of North
America, but even on the Ganges and the Neva. The year in which
Bell’s Comet had come into service Fulton had actually entered into
a contract with one Thomas Lane to introduce steamboats into India,
and on April 12th of that year he wrote to a Russian gentleman, who
was then staying in London, with reference to obtaining an exclusive
contract for twenty years, for establishing a steamboat service
between St. Petersburg and Cronstadt within three years after
obtaining the grant. It is evident from Fulton’s correspondence that
Imperial permission for this was obtained. Fulton, however, died in
the year 1815, and at the time of his death the steamboat The
Emperor of Russia was in course of construction previous to being
transferred to Russian waters. This enterprise was postponed and
subsequently taken up by other contractors. But the same year
(1815) we find Charles Baird engaged in doing what Fulton would
have carried out had he lived. The upper illustration, then, which
faces page 84 represents a drawing of the steamboat Elizabeth.
Originally a barge, she was rebuilt and engined by Baird in 1815 at
St. Petersburg for service on the Neva. The steering arrangement is
not dissimilar to that of some of the Thames sailing barges of to-day,
with the use of the tackle leading from the rudder through the ship’s
quarter to the helm. The reader will doubtless be not a little amused
to notice the brick chimney which stands up in the boat as if rising
from a factory. The engine is hidden away underneath the deck, but
it was of the side-lever type, of which we have already spoken, with
a single cylinder and air-pump. The boiler will be seen placed aft.
The weight of the paddle-wheels was partly supported by the
rectangular frame-work which will be seen stretched across the hull.
The paddle-wheels had each four floats, which were kept level by
means of bevel gear. The other illustration facing page 84 shows
another steamer, which Baird built two years later for passenger
traffic between St. Petersburg and Cronstadt. It will be noticed that,
as in all these early steamboats, the paddle-wheels were placed far
forward towards the bows. In this ship both paddle-wheels were
fitted with six floats, which were driven at fifty revolutions per minute
by means of a side-lever engine that had a large fly-wheel. The
arrangement of this ship’s engines was similar rather to those of the
Comet than of the Clermont. Looking at the lower drawing in this
illustration we can easily see how she was propelled. Amidships is
the boiler, from which steam is conveyed to the cylinder, through
which appears the piston-rod, which in turn connects with the side-
lever, that is placed as low as it can be in the boat. The connecting
rod comes up from the forward end of the side-lever to the crank,
which is attached to the shaft, and the latter, revolving, of course
turns the paddle-wheels.
And here it may not be out of place to say something concerning
the survival of the beam engine. I have already referred on an earlier
page to its introduction and traced its development from
Newcomen’s atmospheric engine. When, in the early days of the
steam engine, its use had been limited to pumping out water from
mines, one connecting rod was employed in pumping and the other
was driven up by the steam in the cylinder. Then, when the engine
was made, not for pumping, but for giving rotatory motion, the
connecting rod which had been in use for pumping was used to give
a rotatory motion, by means of either the sun-and-planet movement
(as in Watt’s patent) or by means of a crank (as in the patent which
his workman stole from him). In America Watt’s beam engines were
imitated very closely, and to-day, as every visitor to New York is
aware, the curious sight is seen of enormous ferry-boats, towering
high above the water, with the beam and connecting rods showing
up through the top of the ship. Now this idea is all very well where
the steamer is concerned only with navigation on rivers and peaceful
waters, but for ocean steaming, where the deck needs to be covered
in from the attacks of the mighty seas, it is out of the question.
Therefore, since it was advisable to retain the beam in some form,
and it could not be allowed to protrude through the deck, the obvious
expedient was adopted of placing it below, but as far down in the
ship as possible. As a general statement we shall not get far wrong if
we state that thus placed, at the bottom, with the rods working
upwards instead of downwards, it was really a case of turning the
engine upside down. Thus arranged it became known as the side-
lever engine, and now, if the reader will look again at the bottom
illustration facing page 84, he will see our meaning. By turning the
illustration round, so that the beam or side-lever is at the top, this
resemblance to the old-fashioned beam engine becomes still more
apparent. Later on we shall be able to show a more complicated
form of the side-lever engine, but for the present this may suffice for
the interest of the non-technical reader. For many years the side-
lever was the recognised form of marine engine, and its advantages
included that of being remarkably steady in its working because its
parts were so nicely balanced. Moreover, it was easy to drive from
the beam the various auxiliary parts, such as the air-pump. It was
also very strong, though both heavy and costly, as it became in the
course of time more complicated.
Although it is true that in Fulton’s Clermont the beam was placed
below the piston-rod, yet that was entirely owing to English influence,
as represented in Boulton and Watt, who had manufactured this
engine, or at any rate a good many of its parts. It is now that the
dividing line comes between the two types, English and American.
“From this primitive form,” says Admiral Preble, in his volume already
quoted, “the two nations diverged in opposite directions—the
Americans navigating rivers, with speed the principal object, kept the
cylinder upon deck and lengthened the stroke of the piston: the
English, on the other hand, having the deep navigation of stormy
seas as their more important object, shortened the cylinder in order
that the piston-rod might work entirely under deck, while Fulton’s
working (walking) beam was retained.” From the engine, in fact,
which Boulton and Watt had constructed at Soho for Fulton, by far
the majority of the engines for the earliest steamboats took their
pattern. And if to the Americans belongs the credit of having so
thoroughly and so quickly developed the steamboat navigation of
large rivers, it is the British, as we shall see shortly, who have been
the pioneers of ocean navigation in steamships.
The upper illustration facing page 90, which has been taken from
a contemporary engraving, is worthy of notice as being the first
steamer actually built in Germany. She represents rather a
retrogression than an advance in the story of the steamship, for she
was following still on those lines which had been in mind when
Miller’s double-hulled ship and the Charlotte Dundas were launched.
This vessel, the Prinzessin Charlotte, was built by John Rubie at
Pichelsdorf in 1816, for service on the Elbe, Havel and Spree. As will
be seen from the illustration, her paddle-wheel was placed
amidships and covered in. She was driven by an engine possessing
14 horse-power and made by J. B. Humphreys. Her long, lanky
smoke-stack is supported by numerous stays, while her double-
rudders, though still preserving the helms as used in contemporary
sailing ships, are moved by means of a steering wheel. Clumsy and
beamy, she is inferior in design to the Comet, and would no doubt
have needed all the help of her twin-rudders to get her round some
of the narrow reaches of the river. In the adoption and employment
of the steering wheel neither the Prinzessin Charlotte nor the
Clermont was the pioneer of this more modern method, its evolution
having come about on this wise: as the tillers became heavier when
the size of ships increased and the pull on them became greater,
some sort of lanyard was first attached to them so as to get a
purchase and divide the strain; otherwise the steersman would not
have been able to control the ship. We see this as far back as the
times of the Egyptian sailing ships. In medieval times and even in the
seventeenth century the big, full-rigged ships were still steered by a
helm in the stern, the pilot shouting down his orders to the
steersmen placed under the poop. Then, in order to counteract the
wild capers which some of these vessels had a tendency to perform
in a breeze, it was an obvious expedient to fit up an arrangement of
blocks and tackles to the tiller. From this came the transition to the
employment of these in connection with a winch, such as had been
used for hoisting up the anchor. This winch was driven by means of
“hand-spikes,” a method that was not conducive to rapid alteration of
the ship’s course. But in the eighteenth century, when ships were
better designed, and many improvements were being introduced, the
handspikes were discarded and the spoked wheel was connected
with the barrel of the winch, placed not ’thwart-ship, but fore-and-aft,
so that not merely could the direction of the ship’s head be altered
more quickly, but a steadier helm could be kept, because it was less
difficult to meet the swervings of the vessel from her proper course.
As everyone knows, this steering-wheel has been improved by many
minor alterations, and ropes have given way to chains and steel
wire: but though steam-steering gear is now so prominent a feature
of the modern steamship, the wheel itself is not yet superseded.
THE “PRINZESSIN CHARLOTTE” (1816).

From a Contemporary Print.
THE “SAVANNAH” (1819).
Already, then, the steamboat had shown herself capable of doing

her work on inland waters, and even for short voyages across
Channel, as well as for coasting within sight of land. Independent of
calms, currents and tides, she was a being of a different kind as
compared with the sailing ship and was carving out for herself an
entirely novel career of usefulness. But the pessimists believed that
here her sphere ended; the long ocean voyages could never be
undertaken except in the sail-carrying ships. However, in the year
1819, the first attempt was made to conquer the North Atlantic by
means of a ship fitted with a steam engine. In the lower illustration
facing page 90 will be seen the Savannah, a full-rigged ship of 350
tons burthen which was built in New York in 1818 as a sailing vessel
pure and simple. That, it will be remembered, was eleven years after
the launching of the Clermont, and during these eventful years there
had been plenty of opportunity for those who wished to obtain proof
of what steam could do for a ship. Whilst the Savannah was still on
the stocks, one Moses Rogers, who had followed the efforts of both
Stevens and Fulton, and had even commanded some of the early
steamboats, suggested to Messrs. Scarborough and Isaacs, of
Savannah, that they should purchase this ship; which eventually they
did. Therefore, after being fitted with her engine, a steam trial trip
was made in March, 1819, round New York Harbour, and a few days
later she left for Savannah under sail. During this voyage of 207
hours she was practically nothing but a sailing ship, for her engine
was only running for four and a half hours. On the 22nd of May she
set forth from Charleston and steamed outside. It will be noticed on
referring to the illustration that there were no paddle-boxes to cover
her wheels, and a remarkable feature of the Savannah was her
ability suddenly to transform her character as a steamship to a
sailing vessel, and vice versa. Within twenty minutes she could take
off her paddle-wheels, and away she could go without any hindrance
to her speed.
So it was, then, after she had brought up outside Charleston.
Unshipping her wheels she got under weigh early in the morning of
May 24th, and arrived off the coast of Ireland at noon of June 17th,
and three days later was off the bar at Liverpool. But this voyage
proved little or nothing of the capabilities of the ocean steamship; for
of the twenty-one days during which she was at sea the Savannah
only used steam for eighty hours, and by the time she had arrived off
Cork she had used up all her fuel. However, having now taken on
board what she needed, she was able to steam up the Mersey with
the aid of her engines alone. From Liverpool she went to the Baltic,
using her engine for about a third of the passage. Thence she
returned to America, having unshipped her paddle-wheels off
Cronstadt, but, after crossing the Atlantic and arriving off the
Savannah river, she adjusted her wheels once more and steamed
home. Shortly afterwards her engines were taken out of her, and she
ended her days as a sailing packet. Although her voyages did
nothing to help forward the ocean steamer, yet she caused some
amazement to the revenue cruiser Kite, which espied her off the
coast of Ireland. Seeing volumes of smoke pouring out from this
“three-sticker,” the Kite’s commander took her for a ship on fire and
chased her for a whole day. The illustration gives a fairly accurate
idea of the ship, though the bow has not been quite correctly given,
and should show the old-fashioned and much modified beak which
survived as a relic of medieval times. It will be noticed that the
distance which separates the main and fore-mast was sufficiently
great to allow of plenty of room for the engine and boiler.
In the meantime the steamship was slowly but surely coming into
prominence and recognition, and the year 1821 was far from
unimportant as showing the practical results which had been
obtained. As proof of the faith which was now placed in steam, the
first steamship company that was ever formed had already been
inaugurated the year before, and in 1821 began running its trading
steamers. This was the now well-known General Steam Navigation
Company, Ltd., whose first steamer, the City of Edinburgh, was built
on the Thames by Messrs. Wigram and Green, whose names will
ever be associated with the fine clippers which in later years they
were destined to turn out from their Blackwall yard. The steamship
City of Edinburgh was launched in March, 1821, for the Edinburgh
trade, and created so much attention that the future William IV. and
Queen Adelaide paid her a visit, and expressed surprise at the
magnificence of the passenger accommodation. The machinery
(which was only of 100 horsepower) was described by the
contemporary press as “extremely powerful.” In June of that year
was also launched the James Watt, of which an illustration is given
from an old water-colour. This vessel was built by Messrs. Wood and
Co., of Port Glasgow, and was referred to by the newspapers of that
time as “the largest vessel ever seen in Great Britain propelled by
steam.” The James Watt, it will be seen, was rigged as a three-
masted schooner, with the typical bow and square stern of the
period. She was of 420 tons, and measured 141 feet 9 inches in
length, 25½ feet wide, and 16½ feet deep. She had a paddle-wheel,
18 feet in diameter, on either side of the hull. These were driven by
engines of the same horsepower as those of the City of Edinburgh,
which had been made by Boulton and Watt. It was in this year also
that the Lightning, a vessel of about 200 tons and 80 horse-power,
gained further confidence for the newer type of vessel, for she was
the first steamship ever used to carry mails.
Before the third decade of the nineteenth century was closed, a
little vessel named the Falcon, of 176 tons, had made a voyage to
India—of course, via the Cape—and the Enterprise, a somewhat
larger craft of 470 tons, had also done the passage from England to
Calcutta; but like the Savannah’s performance, these voyages were
made partly under steam and partly under sail, so that these vessels
may be regarded rather as auxiliary-engined than as steamships
proper. At the same time, the Enterprise was singularly loyal to her
name, for out of the 113 days which were taken on the voyage, she
steamed for 103.
THE “JAMES WATT” (1821).

From a Water-Colour Drawing in the Victoria and Albert Museum.
SIDE-LEVER ENGINES OF THE “RUBY” (1836).
From the Model in the Victoria and Albert Museum.
Let us now pause for a moment to witness some of the changes

which were going on in regard to the machinery for steamships. In
the engines which were installed in the Russian ship shown opposite
page 84 we saw how the beam had become the side-lever, and why
it had been placed in this position in the steamboat. This had
become the customary type for steamships which were still propelled
by paddle-wheels, and the perfected development had been due to
Boulton and Watt, dating from about 1820. Until about 1860 this type
was used most generally, until ocean-going steamers discarded the
paddle-wheel for the screw. It is, therefore, essential that before
proceeding farther we should get well-acquainted with it, and we
shall find that following the lead which had been given them,
especially by the famous Robert Napier, marine engineers began to
build these types, as well for deep-sea ships as for river-going craft.
The illustration here facing, which has been taken from a model in
the South Kensington Museum, represents the regular side-lever
type, the full-sized engines having been made by a Poplar firm in
1836 for the Ruby, which plied between London and Gravesend, a
vessel of 170 tons, and the fastest Thames steamer of that time. On
referring to our illustration, the side-lever will be immediately
recognised in the fore-ground at the bottom. To the left of this are the
two cylinders, side by side. The side-lever is seen to be pivoted at its
centre, whilst at the reader’s left hand the end of this is joined by a
connecting rod. Thus, as the piston-rod is moved upwards or
downwards, so the left-hand half of the side-lever will move. At the
opposite, right-hand, side of the latter the connecting rod will be
observed to be attached to the side-lever, whilst the other end of the
connecting rod drives the crank; the latter, in turn, driving the shaft
on either end of which will be placed a paddle-wheel. In this engine
before us there are two cranks, of which one is seen prominently at
the very top of the picture. Each connecting rod is attached to two
side-levers, one on either side of the cylinder, by means of a cross-
head. Similarly at the piston-rod there is also a cross-head, with a
connecting rod on either side, of which one only is visible. Later on a
modified form of this type of engine was introduced in order to
economise space, for one of the great drawbacks of the side-lever
engine was that it took up an enormous amount of room, which could
ill be spared from that to be devoted to the carrying of cargo or the
accommodation of the passengers. In this modification the cylinders,
instead of being placed side by side, or athwartships, were fore and
aft, the one behind the other.
In 1831, there was built in Quebec, to run between there and
Halifax, a steamer called the Royal William (not to be confused with
a vessel of the same name to which we shall refer presently). The
engines were made by Boulton and Watt, and dispatched across the
Atlantic to Montreal, where they were installed. In 1833, after taking
on board over three hundred tons of coal at Pictou, Nova Scotia, she
started on her journey to the South of England, and arrived off
Cowes, Isle of Wight, after seventeen days, having covered a
distance of 2,500 miles. There is some doubt as to whether she
steamed the whole way, or whether she used her sails for part of the
time. At any rate, she measured 176 feet long, 43 feet 10 inches
wide (including her paddle-boxes), and after calling at Portsmouth,
proceeded to Gravesend, and was afterwards sold to the Spanish
Government.
THE “SIRIUS” (1838).

From a Contemporary Drawing in the Victoria and Albert Museum.
THE “ROYAL WILLIAM” (1838).
By permission of the City of Dublin Steam Packet Co.
We now come to the year 1838, in which a handful of steamers

made history, and showed how uncalled-for had been the ridicule
which the pessimists had cast at the steamship. With this year we
reach the turning-point of the steamship, and from that date we may
trace all those wonderful achievements which are still being added to
year by year. Hitherto no vessel had crossed the Atlantic under
steam power solely. Because of the large amount of fuel
consumption which was a necessary failing of the early steamships,
in proportion to the amount of steam developed, it was denied that it
would ever be financially possible for steamers to run across oceans
as the sailing packets were doing, even if they were capable of
carrying sufficient fuel together with their passengers and cargo. But
deeds were more eloquent than the expounding of theories, and the
first surprise was quickly followed by another, far from inferior. The
first of these epoch-making steamers was the Sirius. She was rigged
as a brig, like many of the contemporary sailing ships which then
carried mails, passengers, and cargo between the Old World and the
New, whose unsavoury characters had earned for them the
nickname of “coffin-brigs.” This Sirius was a comparatively small ship
of 703 tons, and quite small enough to cross the Atlantic in the
weather which is to be found thereon. She measured only 178 feet
along the keel, was 25½ feet wide, her hold was 18¼ feet deep, and
her engines developed 320 horsepower. Built for the service
between London and Cork, she was specially chartered for this
transatlantic trip by the British Queen Steam Navigation Company,
whose own vessel, the British Queen (shown opposite page 102),
was not yet ready, owing to the fact that one of her contractors had
gone bankrupt. With ninety-four passengers on board, the Sirius
steamed away from London and called at Queenstown, where she
coaled. After clearing from the Irish port, she encountered head
winds, and it was only with difficulty that her commander, Lieut. R.
Roberts, R.N., was able to quell a mutiny among the crew, who had
made up their minds that to try and get across the North Atlantic in
such a craft was pure folly. Having been seventeen days out, the
Sirius arrived off New York on April 22nd, and before the end of her
journey had not merely consumed all her coal, at a daily average of
24 tons, but had even to burn some of her spars, so that she had got
across just by the skin of her teeth. But it was her engines which had
got her there and not her sails; the former were of the side-lever type
to which we have just referred.
The next day came in the Great Western, a much larger craft,
that had come out of Bristol three days after the Sirius had started;
and in her we see the prototype of those enormous liners which go
backwards and forwards across the Atlantic to-day with a regularity
that is remarkable. Unlike the little Sirius, the Great Western had
been specially designed for the Atlantic by that engineering genius,
Brunel, who, like his ships and his other works of wonder, was one of
the most remarkable products of the last century. She was built with
the intention of becoming practically an extension of the Great
Western Railway across the Atlantic, and in order to be able to
withstand the terrible battering of the seas, which she would have to
encounter, she was specially strengthened. Here was a vessel of
1,321 tons (gross), with a length of 236 feet over all, with about half
her space taken up with her boilers and engines. Now the strain of
so much dead-weight in so long a ship whose beam was only 35 feet
4 inches, or about one-seventh of her length, had to be thought out
and guarded against with the greatest care. And let us not forget that
at this time vessels were still built of wood, and that, except in a few
instances, iron had not yet been introduced. She was given strong
oak ribs, placed close together, while iron was also used to some
extent in fastening them. The advantage of making an ocean-going
vessel long is that she is less likely to pitch in a sea, and will not dip
twice in the same hollow; and if she is proportionately narrow in
comparison with her length, she will also roll less than a more beamy
craft. But the difficulty, so long as wood was employed, was to get
sufficient longitudinal strength to endure the strains of so long a
span. We shall be able to get some idea of this when we consider
the behaviour of a vessel in a sea. Waves consist, so to speak, of
mountains and valleys. If the waves are short and the vessel is long,
then she may stretch right over some of them; but if the contrary is
the condition, then, while her ’midship portion is supported by the
water, her fore and aft ends are inclined to droop, so that in a very
extreme case she would break in two. At any rate, the tendency is
for the centre of the ship to bend upwards and the unsupported ends
to droop. This is technically called “hogging.” In the reverse
circumstance, when the ends are supported on the tops of two
mountains of waves, whilst the centre of the ship spans,
unsupported, the intervening valley, the tendency is to “sag.” Now
this has to be allowed for in the construction of the ship, and, as
already pointed out in my “Sailing Ships and Their Story,” this was
understood as far back as the times of the Egyptians, who
counteracted such strains as these by means of a longitudinal cable
stretched tightly from one end of the ship to the other. But with the
coming of steamships there was another problem to be taken into
consideration. Engines, boilers, fresh water for the boilers, coal and
so on are serious weights to be placed in one part of the ship. (In the
case of the Great Western, the first three alone weighed 480 tons,
although the gross tonnage of the whole ship was only 1,321.)
Throughout the length of the ship, then, she is subjected not
merely to irregular strains by the peaks and valleys of the waves, but
by the distribution of weights. Her structure has to undergo the
severest possible stresses, and these are different when the ship is
loaded and when she is “light.” If you divide a ship into sections
transversely, as is actually done by the designer, you will find that
some parts are less buoyant than others, no matter whether your
ship is made of wood, iron, or steel. Those sections, for instance,
which contain a steamer’s machinery will have much inferior
buoyancy, and, indeed, were you to sever them from the ship and
seal them up so as to be perfectly water-tight, they would in many
cases sink. Therefore, this irregularity of buoyancy has to be met by
making the more-buoyant sections help to support the less-buoyant.
In actual shipbuilding practice it is customary to regard the greatest
stress to a ship as occurring when she is poised on the crest of a
wave, and it is usual to suppose, in order to safeguard her manner of
construction, that she is poised upon the crest of a wave whose
length from trough to trough is equal to the length of the ship, and
the height of the wave from trough to crest to be one-twentieth of its
length when 300 feet long and below, and one twenty-fifth when
exceeding that length.
We have digressed a little from our immediate subject in order to
put into the mind of the general reader some conception of the
difficulties which Brunel had to encounter when he set to work to
produce such a vessel as the Great Western. That she was built on
sound lines is proved by the service which she rendered to her
owners before she was finally broken up in 1847. On her first return
voyage from New York she took fifteen days, and the Sirius
seventeen. The Great Western had no such trouble with her “coal-
endurance” on her maiden voyage as the Sirius had suffered, for she
had reached New York with one quarter of her coals still
unconsumed, and the obvious conclusion which came to any
reasoning mind was that it certainly paid to build a vessel big enough
to carry plenty of fuel. But the Great Western “paid” in more senses
than this; and at the end of her first year, her directors were able to
announce a dividend of 9 per cent. Thirty-five guineas was the fare
in those days, and the largest number of passengers carried on any
one of her journeys was 152.
THE “GREAT WESTERN” (1838).
By permission of Messrs. Henry Castle & Sons.
PADDLE-WHEEL OF THE “GREAT
WESTERN.”
From the Model in the Victoria and Albert Museum.
Like her contemporaries, the Great Western was fitted with side-
lever engines, built by Maudslay. Steam was generated from four
boilers, and conducted into two cylinders, her daily consumption of
coal being about 33 tons. A model of one of her paddle-wheels,
which were 28 feet 9 inches in diameter, is here illustrated. This type
is known as the “cycloidal” wheel, in which each float, instead of
being made of one solid piece of material, is composed of several
horizontal widths arranged after the manner of steps in a cycloidal
curve, as will be seen by looking at the right-hand of the wheel. It will
be noticed that through the space left between each “step” the water
could penetrate when the wheel was in the sea, but when revolving
out of it, the resistance to the air was diminished because the latter
was allowed to get through. As the paddle came in contact with the
sea, the concussion was lessened, and thus there was not so much
strain on the engines. The Great Western employed the type
introduced by Joshua Field in 1833, but this form was brought in
again by Elijah Galloway two years later.
So far we have seen steamers running from London and from
Bristol to New York. Now we shall see the first steam-vessel crossing
from Liverpool to New York. Facing page 96 is the other Royal
William, which was built in 1838 for the Irish passenger trade
between Liverpool and Kingstown, and owned by the City of Dublin
Steam Packet Company, by whose courtesy this picture is now
reproduced. The Royal William was 3 feet shorter than the Sirius, but
2 feet wider, and with a hold just 6 inches shallower. In July of that
same memorable year, the Royal William made her maiden trip from
Liverpool to New York, having been built and engined at the former
port. In was no doubt a great temptation to emulate what the Sirius
had been the first to perform, especially as the two ships were so
similar in many respects. Outward bound, the Royal William did the
trip in about the same time as the Sirius, though her return journey
occupied about a day and a half less than that of the other vessel.
But these vessels were not big enough, nor seaworthy enough, for
the toil of the Atlantic, and both were soon taken off from this route.
The illustration reproduced is from an engraving after a sketch made
of the Royal William, as seen in the Atlantic on July 14th, 1838, when
in latitude 47.30 N., longitude 30.0 W., on her first voyage to New
York, and the landsman in looking at the waves which the artist has
depicted may find some assistance in reading our previous remarks
on “hogging” and “sagging” in this connection.
THE “BRITISH QUEEN” (1839).
By permission of James Napier, Esq.
THE “BRITANNIA,” THE FIRST ATLANTIC LINER

(1840).
From a Model. By permission of the Cunard Steamship Co.
Finally, we come to the British Queen, which was yet another

vessel to steam across the broad Atlantic, and to show once more
that it was neither good fortune nor the powers of any single vessel
that had conquered the ocean, but the building of the right kind of

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Rhizosphere Engineering Ramesh Chandra Dubey

Fundamental Immunology Seventh Edition

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About the Author ii

3.7 Dendritic Cells 55

7.1 Introduction 110

8.1 Introduction 133

9.1 Introduction 159

10.1 Introduction 178

11.1 Introduction 199

12.1 Introduction 218

13.1 Introduction 245

14.1 Introduction 270

15.1 Introduction 294

16.1 Introduction 310

16.5 The Paradox: Tumors Elude the Immune System 318

Types of Grafts 325

Radioimmunoassay (RIA) 330

● Important terminologies highlighted

●Chapter end questions: 326

Edward Jenner’s pioneering contributions enabled the eradication

THE “PRINZESSIN CHARLOTTE” (1816).

Already, then, the steamboat had shown herself capable of doing

THE “JAMES WATT” (1821).

Let us now pause for a moment to witness some of the changes

THE “SIRIUS” (1838).

We now come to the year 1838, in which a handful of steamers

THE “BRITANNIA,” THE FIRST ATLANTIC LINER

Finally, we come to the British Queen, which was yet another

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