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Principles and
Applications
Contents
Preface to Second Edition
Preface to First Edition
Abbreviations Used in This Book
Greek Letters Used i n This Book
Color Coding for Molecules
Chapter 1: Viruses and Their Importance
1 . 1 Viruses are Ubiquitous on Earth
1.2 Reasons for Studying Viruses
1.3 The Nature of V iruses
1.4 The Remainder of the Book
Learning Outcomes
Sources of Further Information
Chapter 2: Methods Used in Virology
2.1 Introduction to Methods Used in Virology
2.2 Cultivation of Viruses
2.3 Isolation of Viruses
2.4 Centrifugation
2.5 Structural Investigations of Cells and Virions
2.6 Electrophoretic Techniques
2.7 Detection of Viruses and Virus Components
2.8 Infectivity Assays
2. 9 Virus Genetics
2.10 Investigation of ProteinslJ"Protein Interactions
Learning Outcomes
Sources of Further Information
Chapter 3: Virus Structure
3 . 1 Introduction to Virus Structure
3.2 Virus Genomes
3.3 Virus Proteins
3.4 Capsids
3.5 Virion Membranes
3.6 Occlusion Bodies
3.7 Other Virion Components
Learning Outcomes
Sources of Further Information
Chapter 4: Virus Transmission
4. 1 Introduction to Virus Transmission
4.2 Transmission of Plant Viruses
4.3 Transmission of Vertebrate Viruses
4.4 Transmission of Invertebrate Viruses
4.5 Permissive Cells
Learning Outcomes
Sources of Further Information
Chapter 5 : Attachment and Entry of Viruses into Cells
5.1 Overview of Virus Replication
5.2 Animal Viruses
5.3 Bacteriophages
Learning Outcomes
Sources of Further Information
Chapter 6: Transcription, Translation, and Transport
6. 1 Introduction to Transcription, Translation, and Transport
6.2 Transcription of Virus Genomes
6.3 Transcription in Eukaryotes
6.4 Translation in Eukaryotes
6.5 Transport of Molecules in Eukaryotic Cells
6.6 Transcription and Translation in Bacteria
Learning Outcomes
Sources of Further Information
Chapter 7 : Virus Genome Replication
7 . 1 Overview of Virus Genome Replication
7.2 Locations of Virus Genome Replication in Eu karyotic Cells
7 .3 Initiation of Genome Replication
7. 4 Polymerases
7.5 DNA Replication
7 .6 Double-Stranded RNA Replication
7.7 Single-Stranded RNA Replication
7.8 Reverse Transcription
Learning Outcomes
Sources of Further Information
Chapter 8 : Assembly and Exit of Virions from Cells
8 . 1 Introduction to Assembly and Exit of Virions from Cells
8.2 Nucleocapsid Assembly
8.3 Formation of Virion Membranes
8.4 Virion Exit From the Infected Cell
Learning Outcomes
Sources of Further Information
Chapter 9 : Outcomes of Infection for the Host
9 . 1 Introduction to Outcomes of Infection for the Host
9.2 Factors Affecting Outcomes of Infection
9.3 Non-Productive Infections
9.4 Productive Infections
Learning Outcomes
Sources of Further Information
Chapter 10: Classification andl Nomenclature of Viruses
10. 1 Hi story of Virus Classification and Nomenclature
10.2 Modern Virus Classification and Nomenclature
10.3 Baltimore Classification of Viruses
Learning Outcomes
Sources of Further Information
Chapter 11: Herpesviruses (and Other dsDNA Viruses)
1 1 . 1 Introduction to Herpesviruses
1 1 . 2 The Human Herpesviruses
1 1 . 3 The Herpesvirus Virion
11.4 HSV-1 Genome Organization
1 1 . 5 HSV-1 RepI ication
1 1 . 6 Latent Herpesvirus Infection
11.7 Other dsDNA Viruses
Learning Outcomes
Sources of Further Information
Chapter 12: Parvoviruses (and Other ssDNA Viruses)
12.1 Introduction to Parvoviruses
12.2 Examples of Parvoviruses
12.3 Parvovirus Virion
12.4 Parvovirus Replication
12.5 Other ssDNA Vi ruses
Learning Outcomes
Sources of Further Information
Chapter 13: Reoviruses (and Other dsRNA Viruses)
13.l Introduction to Reoviruses
13.2 Rotavirus Virion
13.3 Rotavirus Repl ication
13.4 Other dsRNA Viruses
Learning Outcomes
Sources of Further Information
Chapter 14: Picornaviruses (and Other Plus-Strand RNA Viruses)
14. 1 Introduction to Picornavi ruses
14.2 Some Important Picornaviruses
14.3 Picornavirus Virion
14.4 Picornavirus Replication
14.5 Picornavirus Recombination
14.6 Picornavirus Experimental Systems
14.7 Other Plus-Strand RNA Vi ruses
Learning Outcomes
Sources of Further Information
Chapter 15: Rhabdoviruses (and Other Minus-Strand RNA Viruses)
15. 1 Introduction to Rhabdoviruses
15.2 Some Important Rhabdoviruses
15.3 The Rhabdovirus Virion and Genome Organization
15.4 Rhabdovirus Replication
15.5 Other Minus-Strand RNA Viruses and Viruses with Ambisense Genomes
15.6 Reverse Genetics
Learning Outcomes
Sources of Further Information
Chapter 16: Influenza Virus
16.1 Introduction to Influenza Viruses
16.2 The Influenza Virion
16.3 Influenza A Virus Replication
16.4 Importance of Influenza Viruses
16.5 Reverse Genetics
Learning Outcomes
Sources of Further Information
Chapter 17: Retroviruses
17 . 1 Introduction to Retrovi ruses
17.2 Retrovirus Virion
17.3 Retrovirus Replication
17.4 Examples of Retroviruses
17.5 Retroviruses as Gene Vectors
17 .6 Endogenous Retroviruses
Learning Outcomes
Sources of Further Information
Chapter 18: Human Immunodeficiency Viruses
18.1 Introduction to HIV
18.2 HIV Virion
18.3 HIV Genome
18.4 HIV-1 Replication
18.5 HIV-1 Variability
18.6 Progression of HIV Infection
18.7 Prevention of HIV Transmission
Learning Outcomes
Sources of Further Information
Chapter 19: Hepadnaviruses (and Other Reverse-Tra nscribing DNA Viruses)
19.1 Introduction to Hepadnaviruses
19.2 Importance of HBV
19.3 HBV Virion
19.4 Non-Infectious Particles
19.5 Soluble Virus Protein
19.6 HBV Genome
19.7 HBV Genetic Groups
19.8 HBV Replication
19.9 Prevention and Treatment of HBV Infection
19.10 Other Reverse-Tra nscri bing DNA Viruses
Learn ing Outcomes
Sources of Further Information
Chapter 20: Bacterial Viruses
20.1 Introduction to Bacterial Viruses ( Bacteriophages)
20.2 Single-Stranded RNA Phages
20.3 Double-Stranded RNA Phages
20.4 Single-Stranded DNA Phages
20.5 Double-Stranded DNA Phages
Learning Outcomes
Sources of Further Information
Chapter 2 1 : Origins and Evolution of Viruses
2 1 . 1 Introduction to Origins and Evolution of Viruses
21.2 Origins of Viruses
21.3 Evolution of Viruses
Learning Outcomes
Sources of Further Information
Chapter 22: Emerging Viruses
22.1 Introduction to Emerging Viruses
22.2 Viruses in New Host Species
22.3 Viruses in New Areas
22.4 Viruses in New Host Species and in New Areas
22.5 New viruses
22.6 Recently Discovered Viruses
22. 7 Re-Emerging Viruses
22.8 Virus Surveil lance
22.9 Dealing with Outbreaks
Learning Outcomes
Sources of Further Information
Chapter 23: Viruses and Cancer
23. 1 Introduction to Viruses and Cancer
23.2 Papillomavi rus-Linked Cancers
23.3 Polyomavirus-Linked Cancers
23.4 Epsteinsl.>"Barr Virus-Linked Cancers
23.5 Kaposisl.>™s Sarcoma
23.6 Adult T Cell Leukemia
23.7 Hepatocellular Carcinoma
23.8 Virus-Associated Cancers in Animals
23.9 Cell Lines Derived From Virus-Associated Cancers
23. 1 0 How Do Viruses Cause Cancer?
23. 1 1 Prevention of Virus-Induced Cancers
Learning Outcomes
Sources of Further Information
Chapter 24: Survival of Infectivity
24.1 Preservation of Virus Infectivity
24.2 Destruction of Virus Infectivity
24.3 Inactivation Targets in Virions
24.4 Inactivation Kinetics
24. 5 Agents that Inactivate Virus Infectivity
Learning Outcomes
Sources of Further Information
Chapter 25: Virus Vaccines
25. 1 Introduction to Virus Vaccines
25.2 Live Attenuated Virus Vaccines
25.3 Inactivated Virus Vaccines
25.4 Virion Subunit Vaccines
25.5 Live Recombinant Virus Vaccines
25.6 Mass-Production of Viruses for Vaccines
25.7 Virus-Like Particles
25.8 Synthetic Peptide Vaccines
25.9 DNA Vaccines
25 .10 Storage and Transport of Vaccines
Learning Outcomes
Sources of Further Information
Chapter 26: Anti-viral Drugs
26.1 Introduction to Anti-viral Drugs
26.2 Development of Anti-viral Drugs
26.3 Examples of Anti-viral Drugs
26.4 Drug Resistance
26.5 Anti-vi ra l Drug Research
Lea rn i ng Outcomes
Sou rces of Fu rther Information
Chapter 27: Pri ons
27 . l Introd uction to Prions
27.2 Tra nsm issi ble Spongiform Encephal opathies
27 .3 The Nature of Prions
27.4 Pri on Di seases
27.5 Pri on Stra i ns
27 . 6 Pri on Tra nsm i ssion
27. 7 The Protei n-Only Hypothesis
Lea rn i ng Outcomes
Sou rces of Fu rther Informati on
Vi rologistss1> Vocabulary
TM
Index

VIROLOGY
PRINCIPLES AND APPLICATIONS

John B. Carter
and
Venelia A. Saunders

[S ormRG]
KATI TPB I ET I h33

y
This edition first published 2013 by John Wiley & Sons Ltd
Copyright © 2013, 2007 John Wiley & Sons Ltd
Cover image: Glass Sculpture of HIV virus courtesy of Wellcome Images
All effo rt has been made to trace and acknowledge ownership of copyright. The
publisher would be glad to hear from any copyright holders whom it has not been
possible to contact.
Registered office
John Wiley & Sons Ltd, The Atrium, Southern Gate, Chichester, West Sussex, P019
8SQ, United Kingdom
For details of our global editorial offices, for customer services, and for information
about how to apply for permission to reuse the copyright material in this book
please see our website at www.wiley.com.
The rights of John Carter and Venetia Saunders to be identified as the authors of
this work has been asserted in accordance with the U K Copyright, Designs and
Patents Act 1988.
All rights reserved. No part of this publication may be reprod uced, stored in a
retrieval syste m , or transm itted, in any form o r by any means, electronic,
mechanical, photocopying, recording or otherwise, except as permitted by the UK
Copyright, Designs and Patents Act 1988, without the prior permission of the
publisher.
Wiley publishes in a variety of print and electronic formats and by print-on-demand.
Some material included with standard print versions of this book may not be
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Designations used by companies to distinguish their products are often claimed as
trademarks. All brand names and product names used in this book are trade
names, service marks, trademarks or registered trademarks of their respective
owners. The publisher is not associated with any product or vendor mentioned in
this book. This p u blication is designed to provide accurate and authoritative
information in regard to the subject matter covered . It is sold on the understanding
that the publisher is not engaged in rendering professional services. If professional
advice or other expert assistance is req u i red, the services of a com petent
professiional should be sought.
Library of Congress Catalog ing-in-Pu blication Data
Carter, John B., 1944-
Virology : principles and applications / John Carter, Venetia Saunders. - 2nd ed.
p.; cm.
Includes bibliographical references and index.
ISBN 978-1-119-99 143-4 (cloth) - ISBN 978- 1-1 19-99142-7 (pbk.)
I. Sau nders, Veneti a A . , 1949- II. Ti tl e.
[DN LM : 1 . Vi ruses. 2. Vi rus Diseases . QW 160]
6 1 6 . 9 ' 1 0 1-dc23
201 204 1 238
ISBN 978- 1 - 1 186-2979-6 ( ebk)
ISBN 978- 1 - 1 186-2977-2 (ebk)
ISBN 978- 1 - 1 186-2976-5 (ebk)
ISBN 978- 1 - 1 1 99-9 14 2-7 (p bk. )
ISBN 978- 1 - 1 1 99-9 143-4 ( h bk)
A cata l og ue record for this book i s available from the Briti sh Li bra ry
Seni or Com missi oni ng Ed itor: Andrew M cAleer
Assistant Ed itor : Kathari ne Earwaker
Ma rketi ng M anag ers : Fran Hunt and Jo U nderwood
Project Editor: J u l iet Booker
To Myra, Robert, Jon, and Mark
Preface to Second Ed iti on
Vi rology conti nues to be a n exciti ng subject and to develop at a ra pid pace. The
i n tro d u cti o n of new la borato ry tec h n iq u e s a nd th e conti n u ed a p p l i cati on of
esta bl ished techni ques are prod uci n g a wealth of new i nformation . There has been
an explosion in the publication of vi rology papers reporting the d i scovery of new
viruses and providi ng deeper i nsights i nto many facets of the subject. We have
tried to reflect these developments i n the second edition, which contai ns m uch new
materi a l , i ncl u d i ng a n add itional chapter (on i nfl u enza vi rus ) . In a n attem pt to
i m prove cl a rity, sections of the text have been rewritten .
In the d i a g ra ms, many of which have been red rawn for i ncl usion i n this ed ition,
there i s a sta ndard color code to d ifferentiate va rious types of n uclei c acid and
protei n molecu les. Please note that, in the interests of cla rity, there have been
some mod ifications to the color code used in the fi rst ed iti on. There is a key to the
c o l o r c o d e o n p a g e x x i x . I n a d d i t i o n , t h e b o o k h a s a w e b s i te
( w w w . w i l e y . co m / co l l eg e/ca rte r) w ith a cco m p a n y i n g tea ch i n g a n d l e a r n i n g
resources, incl ud i ng ani mations of vi rus replication cycles.
We are grateful to the many people who p rovided feed back on the fi rst edition a nd
to those who made suggestions for the second edition . We have i ncorporated many
of the suggestions for new material, thoug h space constrai nts prevented us fro m
i ncorporati ng them a l l . Aga i n , we would be g rateful to receive feed back.
We thank the team at J ohn Wi ley & Sons Ltd for a l l thei r help with this new ed ition
and to our fa m i l ies for their conti nuing support.
John B. Carters€-
John . Ca rter147@ntlworld . com
Venetia A. Sau nderss€-
 s€-Authors are now reti red; both were previously at School of Pharmacy &
Biomolecular Sciences, Liverpool John Moores University, Byrom Street, Liverpool,
L3 3AF, UK.
Preface to First Edition
Virology is a fascinating and rapidly developing subject, and is worthy of study
purely because viruses are interesting! Furthermore, virology is a branch of science
that is of immense relevance to mankind for a host of reasons, not least of which
are the threats to human health caused by viruses, such as HIV, hepatitis B virus,
papillomaviruses, measles, and influenza vi ruses, to mention just a few. There is a
continuing need for trained virologists and it is hoped that this book will play a
small role in helping to fulfill that need. To a large extent the material in the book is
based on virology taught at Uverpool John Moores University.
This is not a textbook of fundamental virology, medical virology, veterinary virology,
plant virology or of bacteriophages, but a bit of each of these! The general pattern
of the book is that principles of virology are covered earlier and applications are
covered later. There is no strict demarcation between the two, however, so the
reader may be made aware of important applications while principles are being
introduced.
The first 10 chapters cover basic aspects of virology. A chapter on methods used in
virology comes early in the book, but could be skimmed to gain an overview of its
contents and thereafter used for reference. There is one chapter on each of the
seven Baltimore classes, concentrating mainly on animal viruses. There is a chapter
devoted entirely to HIV and an extended chapter on phages, reflecting the renewed
interest i n their biology and applications. After a chapter on origins and evolution of
viru ses, there follow five chapters covering various aspects of applied virology,
including vaccines and antiviral drugs. The final chapter is on prions, which are not
viruses but are often considered along with the viruses.
Each chapter starts with s"Dt-bAt a glance,s"DK a brief summary with the dual aim of
giving a flavor of what is coming up and providing a revision aid. Each chapter ends
with a list of learning outcomes and a guide to further reading in books and
journals. The references are mainly from the twenty-first century, but there is a
selection of important papers from the last century.
The book has a web site (www.wiley.com/go/carter) where you can find:
• many references additional to those in the book;

• links to the journal references (to the full text where this is freely available,

otherwise to the abstract);

• links to virology web sites;

• self-assessment questions and answers for each chapter, to reinforce and

extend concepts developed in the book.

A key feature of our book is a standard color code to differentiate various types of
nucleic acid and protein molecules in the diagrams. The color code is explained in
the following pages. It is appreciated that color coding may be of limited value to
individuals who have difficulty i n d ifferentiating colors, so we have also labeled
many of the molecules.
A number of virus replication cycles are described and the reader should be aware
that these are models based on evidence to date; the models may have to be
modified in the light of future evidence. We present the virus replication cycles as
fitting within a general framework of seven steps:
1. Attachment of a virion to a cell
2. Entry into the cell
3. Transcription of virus genes into mRNAs
4. Translation of virus mRNAs into virus proteins
5. Genome replication
6. Assembly of the virus proteins and genomes into virions
7. Exit of the virions from the cell.
We hope that this helps in appreciating how virus replication fits into a general
pattern, and i n comparing the replication cycles of different types of virus. For
some groups of viruses the framework has to be modified, and we make clear
when that is the case.
If you come across an unfam iliar term please consult the Virolog istss"f> ™
Vocabulary at the back of the book. This glossary i111cludes not only virology-specific
terms, but also a selection of terms from cel l biology, m olecular biology,
immunology, and medicine.
A list of abbreviations that are used throughout this book appears on the following
pages.
We wish to thank the many people who have made the production of this book
possible. We thanl< all those who supplied images and those who gave permission
for the use of their images; we are especially g rateful to David Bhella, Tom
Goddard, Kathryn Newton, and Jean-Yves Sgro. Thanks also to Robert Carter for
assistance with images. We acknowledge the contributions of the many students
who have acted as guinea pigs for our teaching materials and who have provided
us with feedback. Grateful thanks also to those who reviewed material for the book
and provided valuable feedback. We are sorry that we were unable to include all
the topics suggested, but if we had done so the book would have run to several
volumes! Many thanks to Rachael Ballard and all at John Wiley & Sons Ltd who
helped the book come to fruition. Finally, thanks to our families for their support
and for their patience during those many hours we spent ensconced in the study.
We hope you find the book useful and we would be interested to hear what you
think of it. We have tried to ensure that there are no errors, but it is probable that
some h.ave slipped through; if you come across any errors please inform us.
John B. Carter
John.Carter! [email protected]
Venetia A. Sau nders
School of Pharmacy & Biomolecular Sciences, Liverpool John Moores Un iversity,
Byrom Street, Liverpool, L3 3AF, UK
Abbreviations Used in This Book
(+) DNA plus strand (positive strand) DNA
( - ) DNA minus strand (negative strand) DNA
(+) RNA plus strand (positive strand) RNA
( - ) RNA minus strand (negative strand) RNA
A adenine
ADP adenosine diphosphate
AIDS acquired immune deficiency syndrome
AP-1 activator proteii n 1
ATP adenosine triphosphate
b base(s)
BL Burkitt's lymphoma
bp base pair(s)
BSE bovine spongiform encephalitis
c cytosine
C term inus carboxy terminus
cccDNA cova lently closed circular DNA
CD cluster of differentiation
cDNA copy DNA
OD Creutzfeldt-Jakob disease
cos cohesive end
CP coat protein
CPE cytopathic effect
DIP defective interfering particle
DNA deoxyribose nucleic acid
ds double-stranded
DTR direct terminal repeat
 XI

0
Fig ure 1 . 2 Transm ission el ectron microsco pe. Th i s is a microscope i n wh ich the
i mage is formed by electrons transmitted through the speci men .
Sou rce : Photograph courtesy of J EOL.

The u n its i n which vi rions are normally measu red are nanometers ( 1 n m = 1 0 s€'9
m ) . Although vi rions are very small , their d i mensions cover a large range. Amongst
the smal lest are parvovi ruses ( Figure 1 . 3), with diameters about 20 n m , while the
megav i rus and m i m ivirus ( m icrobe-m i m icki n g virus), isolated from ame bae, are
amongst the largest.
Fig ure 1 . 3 Vi rions of m i m ivi rus, one of the largest vi ruses, and a parvovi rus, one of
the smal lest vi ruses.
Sou rce : El ectron m icrograph of m i m ivi rus from Cl averie et al . ( 2009) J ou rnal of
Invertebrate Pathology, 1 0 1 , 172. Reproduced by perm ission of Elsevi er and the
authors. Electron m icrog raph of parvov1i ru s by perm issi on of Professor M . Stewart
McN ulty and The Ag ri- Food and Biosci ences Institute .
Another random document with
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Newgate or Bridewell. It will be a great comfort for our brother
Drayton to hear that he is not chargeable to the Society during his
imprisonment. Truly the Lord hath blessed him in his deed, for in
taking these children he hath brought a blessing to his household.
Doubtless the Lord hath spoken by him to me concerning the
manner He would have me use that which He hath given me, for of
all the plans that hath been devised, this for the founding of a new
Quaker colony across the seas cloth commend itself to me as the
wisest and most useful."

"Verily, it was through these children that the Spirit gave him that
wisdom," said the old man. "He hath come to me sometimes, and
told me how sorely his heart ached for the little ones, and how
hopeless it seemed that they could ever dwell together again. It
would cost Westland years of labour even to send for his wife, and
he could never hope to be able to pay for all his children to go to the
plantations to him. Yet it was the only earthly hope our martyr-brother
cherished, and each time that Friend Drayton went to see him, his
talk would be of the home he would make for his wife and children
across the seas."

"Yea, and verily his hopes shall be fulfilled," said Sir William
fervently. "We will have a free colony where no man shall dare to
say, 'Ye cannot serve God after this fashion,' but where we may lift
up our voices in prayer and praise, none daring to make us afraid."

Audrey thought that such a place might be a little heaven below to

some people, but she was not so sure that Bessie's father would be
happy there, for if there were only Quakers to live in the colony, there
would be no scope for his preaching-power, and he would have to do
as Bessie had done, turn the energy to a more practical account;
and the fervour once displayed in preaching to sinners who would
not hear, might be used in some way for the help and development
of the Friends dependent upon each other for all the comfort and joy
of life.

But seeing what Bessie had done in the way of practical work that
lay nearest to her hand, there was little doubt but that her father
would do the same; for some such thoughts as these had arisen in
the mind of Sir William Penn when the plan was first proposed to
him, and that was why he felt so much pleasure in hearing about
Bessie and the homely work she had undertaken. Out there in his
new colony there would be plenty of homely work for everyone who
would do it, and those who could not stoop to that would be of little
use to themselves or others, and therefore had better stay in
England until they were wiser, or the king grew tired of fining and
imprisoning Quakers. This was not likely to happen very soon,
seeing that these people were a convenient scapegoat for the
gradual curtailment of civil and religious liberty, which was slowly but
surely being effected now in the new laws that were made and put
into force so rigorously.

CHAPTER VIII.
CONCLUSION.

THE six months' imprisonment to which Master Drayton the hatter

had been condemned came to an end at last, but not before his
health had become so greatly impaired by the close confinement and
impure air of the prison that the doctor greatly feared he would never
be strong again, more especially as he would now be known as a
Quaker, and consequently watched and harassed by the authorities
upon the smallest provocation.

The thought of this and the sight of her husband's pale worn face
soon overcame Dame Drayton's reluctance to give up her home and
friends in England, and join the band of Quakers who were soon to
sail for New Jersey in His Majesty's plantations of America.

This year 1677 was likely to be one of blessed memory in the history
of the Society of Friends, for Sir William Penn had carried into
practical effect the dream of Master Drayton, and had spent a portion
of his wealth in the purchase of land upon which his poor persecuted
friends could settle, and worship God according to the dictates of
their conscience, none daring to make them afraid.

Bessie's father had been despatched with a party of convicts to the

older settlement of Massachusetts; but as Quakers were persecuted
almost as bitterly at Boston as in London, an agent had been sent by
Sir William to buy the prisoner at the auction of the convicts, which
would take place as soon as the vessel arrived.

Westland was a more robust man than his friend the hatter, and,
thanks to the care of Friends outside the prison, he had not suffered
so severely as the London tradesman.

The lax rules of the prison had given Westland an opportunity of

preaching the gospel to the prisoners confined with him, and though
many mocked and jeered at his warnings, a few were impressed with
the earnestness of his faith; and this was so great a comfort to his
ardent soul that he forgot the discomfort of his surroundings in the
joy of knowing he had been the means of awakening some souls
from the night and sleep of sin, to seek the Saviour who could give to
them a new and better life.
As it had been decreed that his wife must remain in Bridewell until he
could earn the money to pay her passage to the plantations, it was at
first feared that she would have to remain in prison for a much longer
time than her husband; but although Sir William Penn was known to
be an obstinate Quaker, many about the Court who had known his
father were willing to do him a favour, in the hope of drawing him
back to what they deemed was his rightful position in society; and by
the interest of some of these it was at last arranged that Bessie's
mother should be released when the party of Friends were ready to
sail from Gravesend.

Bessie was allowed to go and see her father just before he was
taken from Newgate; and now to hear at last that her mother would
be released to go with them to the new strange home across the
seas was almost too much joy for the poor girl.

"We shall see her, Dorothy! we shall see her! thee and me; and we
shall not be afraid of people knowing we are Quakers. Verily, God
hath been good in giving us such a friend as Sir William Penn, who
is indeed our champion and protector."

But Audrey was by no means so delighted as Bessie over the

impending change. The two girls had learned to know and love each
other by this time, for each had been drawn to the other by the
mutual helpfulness that had kept business and household going
during the long illness of Dame Drayton and the imprisonment of her
husband.

It had seemed impossible at first that pretty, fashionable Audrey

Lowe, whose father lived by ministering at a steeple-house, and the
stern, uncompromising Bessie Westland could ever be friends, in the
closest sense of that word. But circumstances had thrown them so
closely together the last few months, that they had learned to look
below the surface they each so much disliked in the other, and there
they could recognise the true spirit of Christ Jesus, who came not to
be ministered unto, but to minister.
In spite of her mother's chafings and warnings lest their fashionable
friends should find out that they were related to Quakers, Audrey
Lowe insisted upon spending the greater part of her time at Soper
Lane during her aunt's illness; so that the girls were necessarily
thrown together a good deal, and thus had learned to know and
appreciate the selflessness each displayed in working for Dame
Drayton and her family. When at last the day of parting came, it was
not such an unmingled joy to Bessie as she had anticipated, for her
heart clung to this friend, who was so like and yet so unlike herself in
all externals of character and surroundings.

"I shall never be able to say a word against steeple-houses and the
people who go there, after knowing thee, Audrey," she said, the last
evening they spent together.

They were sitting on a box in the keeping-room, waiting for the

waggon to come and fetch the last of their goods to the wherry,
which would carry them to the schooner chartered by Sir William
Penn to convey them to their new home. The girls sat hand in hand,
their hearts too full to say much, until Bessie spoke about the
steeple-houses.

"I am glad," said Audrey in a whisper, "for I never liked to hear thee
speak of what I loved and reverenced with such contempt. I cannot
understand how thee can do it, when it is God's house of prayer."

But this was treading on dangerous ground, and had been the most
thorny subject of discussion between the two girls; so Audrey
hastened to add—

"I have learned to understand what a real Quaker is from knowing

thee, Bessie; and I shall always try to help them if I can, and they
need my help."

"Ah, and I fear there will be many poor Friends left behind here in
London who will need all the help man can give them," answered
Bessie. "For we cannot all go in this ship to another land; and Friend
William Penn says it would not be good for us or for England to carry
all the Quakers away. We have had our share of the battle, and
fought for the truth and for liberty of conscience, as God
strengthened us to do. Now He will strengthen others to take our
places, while we go to plant the truth in other lands. Although Friend
William Penn hath been imprisoned for the truth again and again, he
will not come with us now, but stay and fight the battle of religious
liberty here; for it can only grow and become strong through fighting
and struggling—hard as it may be for us who suffer."

"Oh, Bessie, I cannot bear to think there should be all this fighting
about it," said Audrey, in a pained tone.

"It hurts thee only to think of it," said Bessie, "and therefore God hath
not called thee to this work, but to be a comforter of those who
suffer, and help to make them strong and gentle. Thou art tender and
loving and pitiful. I thought scornfully of these things once; but since I
have known thee, I have learned to see that God hath work for all in
His world. For it is His world, Audrey, in spite of the sin and pain and
trouble that wicked people make in it. Now I want to fight this
wickedness, and so does my father. But it may be God hath other
methods, only I have not learned them. But I am glad—oh, so glad!
—that God hath called my mother and father, and all of us, out of the
fight for a little while—or, at least, this sort of fighting," added Bessie.

"The fight can never be over, while we have our own sin and
selfishness to struggle against," said Audrey quickly.

"I know. I have learned that since I have been here," replied Bessie.
"There was not time to think of much besides the other sort of
fighting before. We needed all our courage to be faithful and true,
and preach the gospel to every creature, as the Lord Jesus
commanded; but since I have been here, dwelling in safety and
comfort, such as I never knew before, I have learned there is another
battle to fight, and other victories to be won, and I have been trying
to do this as well."

"I know, Bessie," whispered Audrey, "I know it has not been easy for
you to do just the everyday work that was so important to aunt and
uncle. You are Brave Bessie Westland—the bravest girl I ever knew,
especially in what you have done for aunt and all of us here."

They were interrupted at this point, for the box on which they were
seated was wanted, and there was no further opportunity of talking.

At daybreak they were going by water to Limehouse Hole, where a

wherry was to be in readiness to convey them to Gravesend. The
whole party who were going were Quakers, many of them in broken
health from imprisonment in unwholesome gaols. Some were
bringing all their household goods, as Master Drayton was doing;
while others, like Bessie's mother, possessed but the few rags they
wore when leaving prison. Most of them came from London and its
neighbourhood; but a few were brought from neighbouring gaols, the
authorities giving them up to save, the expense of transporting them
as slaves to the plantations.

Audrey and her mother bade the Draytons farewell the night before
they started. It was hard for the sisters to part after this short
reunion, for they too had begun to understand each other better than
they had done before, and whatever their differences of opinion
might be, they were heartily at one in desiring that religious liberty
should be the right of everybody, whatever name they might be
called by; for, as Dame Lowe remarked, there were more silent
martyrs in any cause than the world dreamed of; and, as Audrey
added, there were not many like Brave Bessie Westland.

So the tears of parting had all been shed when the sun rose the next
morning, and if they were not all as happy as Bessie herself and her
two sisters, it was a calm and hopeful party of men and women who
went on board the wherry at Limehouse Hole, and though most of
them were being forcibly driven from their native land, they could yet
look forward to the new home they were going to make in the
unknown world beyond the seas. To many of the more timid of the
company, seated among the baskets and bundles on board the
wherry, the voyage, with its unknown perils, was the most fearful part
of the trial, and if they could not have rested upon the arm of their
Father in heaven, they would scarcely have braved its dangers even
to escape persecution. But almost all among them had a nearer and
dearer self in husband, wife, or children, to think of, and for their
sakes the timid became brave, for the time at least, so that when the
schooner was reached, where the prisoners had been already
placed under the care of the captain, the party of Friends in the
wherry were able to meet them with cheerful, hopeful words and
greetings.

To Bessie and her sisters it was a moment of great joy, although a

second look at the dear mother showed that the months spent in
prison had left cruel marks upon her. The hair, so dark when she
went away, was now quite white, they saw, as they looked more
closely under the hood that covered her head. She was better
clothed than most of those who had been brought from different
gaols, for Dame Lowe and Audrey had made her a homely but useful
outfit for the voyage, and some of the things had been taken by
Friends to the prison the day before. For this thoughtful kindness
Dame Westland was deeply grateful, since the rags she had been
wearing would have been a pain and grief to Bessie, she knew, and
to be able to meet her children decently clad was a great comfort.

Truly the passengers going on this voyage were of all sorts and
conditions of men; but they were linked in the bonds of love to God,
and the truth declared by the Lord Jesus Christ, and for this they had
all suffered in mind, body, or estate, some being beggared, some
maimed, some broken in health and hope alike, but all brave, true
friends and brethren, ready to help each other and bear each other's
burdens.

By the help of the same benefactor, under the guidance of God, they
had been brought together to make a new home in a new country,
and they resolved that, so far as it was possible, religious as well as
civil liberty, should be the charter of the new homestead they were
going to set up in New Jersey. It was to be a home and refuge for the
persecuted Society of Friends. Sir William Penn had bought it, and
they were to establish the faith of God upon it.
Later, perhaps, if the persecution of their people in England did not
cease, he would endeavour to secure a larger territory in liquidation
of a debt owing by the king, for money advanced by his father the
admiral. Whether these larger plans would ever come to anything,
the present band of pilgrims did not know; but, of course, it would
largely depend upon the success of this venture, so every man and
woman of the party felt that it would depend upon them whether or
not this larger refuge could be founded, and all with one accord, who
had heard the story, resolved to follow the example of Brave Bessie
Westland.

MORRISON AND GIBB, PRINTERS, EDINBURGH.

*** END OF THE PROJECT GUTENBERG EBOOK BRAVE BESSIE
WESTLAND ***

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