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Viruses: A Very Short Introduction 3rd

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Viruses: A Very Short Introduction
VERY SHORT INTRODUCTIONS are for anyone wanting a stimulating and
accessible way into a new subject. They are written by experts, and have
been translated into more than 45 different languages.
The series began in 1995, and now covers a wide variety of topics in every
discipline. The VSI library currently contains over 700 volumes—a Very Short
Introduction to everything from Psychology and Philosophy of Science to
American History and Relativity—and continues to grow in every subject
area.

Very Short Introductions available now:

ABOLITIONISM Richard S. Newman


THE ABRAHAMIC RELIGIONS Charles L. Cohen
ACCOUNTING Christopher Nobes
ADOLESCENCE Peter K. Smith
THEODOR W. ADORNO Andrew Bowie
ADVERTISING Winston Fletcher
AERIAL WARFARE Frank Ledwidge
AESTHETICS Bence Nanay
AFRICAN AMERICAN RELIGION Eddie S. Glaude Jr
AFRICAN HISTORY John Parker and Richard Rathbone
AFRICAN POLITICS Ian Taylor
AFRICAN RELIGIONS Jacob K. Olupona
AGEING Nancy A. Pachana
AGNOSTICISM Robin Le Poidevin
AGRICULTURE Paul Brassley and Richard Soffe
ALEXANDER THE GREAT Hugh Bowden
ALGEBRA Peter M. Higgins
AMERICAN BUSINESS HISTORY Walter A. Friedman
AMERICAN CULTURAL HISTORY Eric Avila
AMERICAN FOREIGN RELATIONS Andrew Preston
AMERICAN HISTORY Paul S. Boyer
AMERICAN IMMIGRATION David A. Gerber
AMERICAN INTELLECTUAL HISTORY Jennifer Ratner-Rosenhagen
THE AMERICAN JUDICIAL SYSTEM Charles L. Zelden
AMERICAN LEGAL HISTORY G. Edward White
AMERICAN MILITARY HISTORY Joseph T. Glatthaar
AMERICAN NAVAL HISTORY Craig L. Symonds
AMERICAN POETRY David Caplan
AMERICAN POLITICAL HISTORY Donald Critchlow
AMERICAN POLITICAL PARTIES AND ELECTIONS L. Sandy Maisel
AMERICAN POLITICS Richard M. Valelly
THE AMERICAN PRESIDENCY Charles O. Jones
THE AMERICAN REVOLUTION Robert J. Allison
AMERICAN SLAVERY Heather Andrea Williams
THE AMERICAN SOUTH Charles Reagan Wilson
THE AMERICAN WEST Stephen Aron
AMERICAN WOMEN’S HISTORY Susan Ware
AMPHIBIANS T. S. Kemp
ANAESTHESIA Aidan O’Donnell
ANALYTIC PHILOSOPHY Michael Beaney
ANARCHISM Alex Prichard
ANCIENT ASSYRIA Karen Radner
ANCIENT EGYPT Ian Shaw
ANCIENT EGYPTIAN ART AND ARCHITECTURE Christina Riggs
ANCIENT GREECE Paul Cartledge
THE ANCIENT NEAR EAST Amanda H. Podany
ANCIENT PHILOSOPHY Julia Annas
ANCIENT WARFARE Harry Sidebottom
ANGELS David Albert Jones
ANGLICANISM Mark Chapman
THE ANGLO-SAXON AGE John Blair
ANIMAL BEHAVIOUR Tristram D. Wyatt
THE ANIMAL KINGDOM Peter Holland
ANIMAL RIGHTS David DeGrazia
THE ANTARCTIC Klaus Dodds
ANTHROPOCENE Erle C. Ellis
ANTISEMITISM Steven Beller
ANXIETY Daniel Freeman and Jason Freeman
THE APOCRYPHAL GOSPELS Paul Foster
APPLIED MATHEMATICS Alain Goriely
THOMAS AQUINAS Fergus Kerr
ARBITRATION Thomas Schultz and Thomas Grant
ARCHAEOLOGY Paul Bahn
ARCHITECTURE Andrew Ballantyne
THE ARCTIC Klaus Dodds and Jamie Woodward
ARISTOCRACY William Doyle
ARISTOTLE Jonathan Barnes
ART HISTORY Dana Arnold
ART THEORY Cynthia Freeland
ARTIFICIAL INTELLIGENCE Margaret A. Boden
ASIAN AMERICAN HISTORY Madeline Y. Hsu
ASTROBIOLOGY David C. Catling
ASTROPHYSICS James Binney
ATHEISM Julian Baggini
THE ATMOSPHERE Paul I. Palmer
AUGUSTINE Henry Chadwick
JANE AUSTEN Tom Keymer
AUSTRALIA Kenneth Morgan
AUTISM Uta Frith
AUTOBIOGRAPHY Laura Marcus
THE AVANT GARDE David Cottington
THE AZTECS Davíd Carrasco
BABYLONIA Trevor Bryce
BACTERIA Sebastian G. B. Amyes
BANKING John Goddard and John O. S. Wilson
BARTHES Jonathan Culler
THE BEATS David Sterritt
BEAUTY Roger Scruton
LUDWIG VAN BEETHOVEN Mark Evan Bonds
BEHAVIOURAL ECONOMICS Michelle Baddeley
BESTSELLERS John Sutherland
THE BIBLE John Riches
BIBLICAL ARCHAEOLOGY Eric H. Cline
BIG DATA Dawn E. Holmes
BIOCHEMISTRY Mark Lorch
BIOGEOGRAPHY Mark V. Lomolino
BIOGRAPHY Hermione Lee
BIOMETRICS Michael Fairhurst
ELIZABETH BISHOP Jonathan F. S. Post
BLACK HOLES Katherine Blundell
BLASPHEMY Yvonne Sherwood
BLOOD Chris Cooper
THE BLUES Elijah Wald
THE BODY Chris Shilling
NIELS BOHR J. L. Heilbron
THE BOOK OF COMMON PRAYER Brian Cummings
THE BOOK OF MORMON Terryl Givens
BORDERS Alexander C. Diener and Joshua Hagen
THE BRAIN Michael O’Shea
BRANDING Robert Jones
THE BRICS Andrew F. Cooper
THE BRITISH CONSTITUTION Martin Loughlin
THE BRITISH EMPIRE Ashley Jackson
BRITISH POLITICS Tony Wright
BUDDHA Michael Carrithers
BUDDHISM Damien Keown
BUDDHIST ETHICS Damien Keown
BYZANTIUM Peter Sarris
CALVINISM Jon Balserak
ALBERT CAMUS Oliver Gloag
CANADA Donald Wright
CANCER Nicholas James
CAPITALISM James Fulcher
CATHOLICISM Gerald O’Collins
CAUSATION Stephen Mumford and Rani Lill Anjum
THE CELL Terence Allen and Graham Cowling
THE CELTS Barry Cunliffe
CHAOS Leonard Smith
GEOFFREY CHAUCER David Wallace
CHEMISTRY Peter Atkins
CHILD PSYCHOLOGY Usha Goswami
CHILDREN’S LITERATURE Kimberley Reynolds
CHINESE LITERATURE Sabina Knight
CHOICE THEORY Michael Allingham
CHRISTIAN ART Beth Williamson
CHRISTIAN ETHICS D. Stephen Long
CHRISTIANITY Linda Woodhead
CIRCADIAN RHYTHMS Russell Foster and Leon Kreitzman
CITIZENSHIP Richard Bellamy
CITY PLANNING Carl Abbott
CIVIL ENGINEERING David Muir Wood
CLASSICAL LITERATURE William Allan
CLASSICAL MYTHOLOGY Helen Morales
CLASSICS Mary Beard and John Henderson
CLAUSEWITZ Michael Howard
CLIMATE Mark Maslin
CLIMATE CHANGE Mark Maslin
CLINICAL PSYCHOLOGY Susan Llewelyn and Katie Aafjes-van Doorn
COGNITIVE BEHAVIOURAL THERAPY Freda McManus
COGNITIVE NEUROSCIENCE Richard Passingham
THE COLD WAR Robert J. McMahon
COLONIAL AMERICA Alan Taylor
COLONIAL LATIN AMERICAN LITERATURE Rolena Adorno
COMBINATORICS Robin Wilson
COMEDY Matthew Bevis
COMMUNISM Leslie Holmes
COMPARATIVE LITERATURE Ben Hutchinson
COMPETITION AND ANTITRUST LAW Ariel Ezrachi
COMPLEXITY John H. Holland
THE COMPUTER Darrel Ince
COMPUTER SCIENCE Subrata Dasgupta
CONCENTRATION CAMPS Dan Stone
CONFUCIANISM Daniel K. Gardner
THE CONQUISTADORS Matthew Restall and Felipe Fernández-Armesto
CONSCIENCE Paul Strohm
CONSCIOUSNESS Susan Blackmore
CONTEMPORARY ART Julian Stallabrass
CONTEMPORARY FICTION Robert Eaglestone
CONTINENTAL PHILOSOPHY Simon Critchley
COPERNICUS Owen Gingerich
CORAL REEFS Charles Sheppard
CORPORATE SOCIAL RESPONSIBILITY Jeremy Moon
CORRUPTION Leslie Holmes
COSMOLOGY Peter Coles
COUNTRY MUSIC Richard Carlin
CREATIVITY Vlad Glăveanu
CRIME FICTION Richard Bradford
CRIMINAL JUSTICE Julian V. Roberts
CRIMINOLOGY Tim Newburn
CRITICAL THEORY Stephen Eric Bronner
THE CRUSADES Christopher Tyerman
CRYPTOGRAPHY Fred Piper and Sean Murphy
CRYSTALLOGRAPHY A. M. Glazer
THE CULTURAL REVOLUTION Richard Curt Kraus
DADA AND SURREALISM David Hopkins
DANTE Peter Hainsworth and David Robey
DARWIN Jonathan Howard
THE DEAD SEA SCROLLS Timothy H. Lim
DECADENCE David Weir
DECOLONIZATION Dane Kennedy
DEMENTIA Kathleen Taylor
DEMOCRACY Bernard Crick
DEMOGRAPHY Sarah Harper
DEPRESSION Jan Scott and Mary Jane Tacchi
DERRIDA Simon Glendinning
DESCARTES Tom Sorell
DESERTS Nick Middleton
DESIGN John Heskett
DEVELOPMENT Ian Goldin
DEVELOPMENTAL BIOLOGY Lewis Wolpert
THE DEVIL Darren Oldridge
DIASPORA Kevin Kenny
CHARLES DICKENS Jenny Hartley
DICTIONARIES Lynda Mugglestone
DINOSAURS David Norman
DIPLOMATIC HISTORY Joseph M. Siracusa
DOCUMENTARY FILM Patricia Aufderheide
DREAMING J. Allan Hobson
DRUGS Les Iversen
DRUIDS Barry Cunliffe
DYNASTY Jeroen Duindam
DYSLEXIA Margaret J. Snowling
EARLY MUSIC Thomas Forrest Kelly
THE EARTH Martin Redfern
EARTH SYSTEM SCIENCE Tim Lenton
ECOLOGY Jaboury Ghazoul
ECONOMICS Partha Dasgupta
EDUCATION Gary Thomas
EGYPTIAN MYTH Geraldine Pinch
EIGHTEENTH‑CENTURY BRITAIN Paul Langford
THE ELEMENTS Philip Ball
EMOTION Dylan Evans
EMPIRE Stephen Howe
EMPLOYMENT LAW David Cabrelli
ENERGY SYSTEMS Nick Jenkins
ENGELS Terrell Carver
ENGINEERING David Blockley
THE ENGLISH LANGUAGE Simon Horobin
ENGLISH LITERATURE Jonathan Bate
THE ENLIGHTENMENT John Robertson
ENTREPRENEURSHIP Paul Westhead and Mike Wright
ENVIRONMENTAL ECONOMICS Stephen Smith
ENVIRONMENTAL ETHICS Robin Attfield
ENVIRONMENTAL LAW Elizabeth Fisher
ENVIRONMENTAL POLITICS Andrew Dobson
ENZYMES Paul Engel
EPICUREANISM Catherine Wilson
EPIDEMIOLOGY Rodolfo Saracci
ETHICS Simon Blackburn
ETHNOMUSICOLOGY Timothy Rice
THE ETRUSCANS Christopher Smith
EUGENICS Philippa Levine
THE EUROPEAN UNION Simon Usherwood and John Pinder
EUROPEAN UNION LAW Anthony Arnull
EVANGELICALISM John Stackhouse
EVIL Luke Russell
EVOLUTION Brian and Deborah Charlesworth
EXISTENTIALISM Thomas Flynn
EXPLORATION Stewart A. Weaver
EXTINCTION Paul B. Wignall
THE EYE Michael Land
FAIRY TALE Marina Warner
FAMILY LAW Jonathan Herring
MICHAEL FARADAY Frank A. J. L. James
FASCISM Kevin Passmore
FASHION Rebecca Arnold
FEDERALISM Mark J. Rozell and Clyde Wilcox
FEMINISM Margaret Walters
FILM Michael Wood
FILM MUSIC Kathryn Kalinak
FILM NOIR James Naremore
FIRE Andrew C. Scott
THE FIRST WORLD WAR Michael Howard
FLUID MECHANICS Eric Lauga
FOLK MUSIC Mark Slobin
FOOD John Krebs
FORENSIC PSYCHOLOGY David Canter
FORENSIC SCIENCE Jim Fraser
FORESTS Jaboury Ghazoul
FOSSILS Keith Thomson
FOUCAULT Gary Gutting
THE FOUNDING FATHERS R. B. Bernstein
FRACTALS Kenneth Falconer
FREE SPEECH Nigel Warburton
FREE WILL Thomas Pink
FREEMASONRY Andreas Önnerfors
FRENCH LITERATURE John D. Lyons
FRENCH PHILOSOPHY Stephen Gaukroger and Knox Peden
THE FRENCH REVOLUTION William Doyle
FREUD Anthony Storr
FUNDAMENTALISM Malise Ruthven
FUNGI Nicholas P. Money
THE FUTURE Jennifer M. Gidley
GALAXIES John Gribbin
GALILEO Stillman Drake
GAME THEORY Ken Binmore
GANDHI Bhikhu Parekh
GARDEN HISTORY Gordon Campbell
GENES Jonathan Slack
GENIUS Andrew Robinson
GENOMICS John Archibald
GEOGRAPHY John Matthews and David Herbert
GEOLOGY Jan Zalasiewicz
GEOMETRY Maciej Dunajski
GEOPHYSICS William Lowrie
GEOPOLITICS Klaus Dodds
GERMAN LITERATURE Nicholas Boyle
GERMAN PHILOSOPHY Andrew Bowie
THE GHETTO Bryan Cheyette
GLACIATION David J. A. Evans
GLOBAL CATASTROPHES Bill McGuire
GLOBAL ECONOMIC HISTORY Robert C. Allen
GLOBAL ISLAM Nile Green
GLOBALIZATION Manfred B. Steger
GOD John Bowker
GOETHE Ritchie Robertson
THE GOTHIC Nick Groom
GOVERNANCE Mark Bevir
GRAVITY Timothy Clifton
THE GREAT DEPRESSION AND THE NEW DEAL Eric Rauchway
HABEAS CORPUS Amanda Tyler
HABERMAS James Gordon Finlayson
THE HABSBURG EMPIRE Martyn Rady
HAPPINESS Daniel M. Haybron
THE HARLEM RENAISSANCE Cheryl A. Wall
THE HEBREW BIBLE AS LITERATURE Tod Linafelt
HEGEL Peter Singer
HEIDEGGER Michael Inwood
THE HELLENISTIC AGE Peter Thonemann
HEREDITY John Waller
HERMENEUTICS Jens Zimmermann
HERODOTUS Jennifer T. Roberts
HIEROGLYPHS Penelope Wilson
HINDUISM Kim Knott
HISTORY John H. Arnold
THE HISTORY OF ASTRONOMY Michael Hoskin
THE HISTORY OF CHEMISTRY William H. Brock
THE HISTORY OF CHILDHOOD James Marten
THE HISTORY OF CINEMA Geoffrey Nowell-Smith
THE HISTORY OF COMPUTING Doron Swade
THE HISTORY OF LIFE Michael Benton
THE HISTORY OF MATHEMATICS Jacqueline Stedall
THE HISTORY OF MEDICINE William Bynum
THE HISTORY OF PHYSICS J. L. Heilbron
THE HISTORY OF POLITICAL THOUGHT Richard Whatmore
THE HISTORY OF TIME Leofranc Holford‑Strevens
HIV AND AIDS Alan Whiteside
HOBBES Richard Tuck
HOLLYWOOD Peter Decherney
THE HOLY ROMAN EMPIRE Joachim Whaley
HOME Michael Allen Fox
HOMER Barbara Graziosi
HORMONES Martin Luck
HORROR Darryl Jones
HUMAN ANATOMY Leslie Klenerman
HUMAN EVOLUTION Bernard Wood
HUMAN PHYSIOLOGY Jamie A. Davies
HUMAN RESOURCE MANAGEMENT Adrian Wilkinson
HUMAN RIGHTS Andrew Clapham
HUMANISM Stephen Law
HUME James A. Harris
HUMOUR Noël Carroll
THE ICE AGE Jamie Woodward
IDENTITY Florian Coulmas
IDEOLOGY Michael Freeden
THE IMMUNE SYSTEM Paul Klenerman
INDIAN CINEMA Ashish Rajadhyaksha
INDIAN PHILOSOPHY Sue Hamilton
THE INDUSTRIAL REVOLUTION Robert C. Allen
INFECTIOUS DISEASE Marta L. Wayne and Benjamin M. Bolker
INFINITY Ian Stewart
INFORMATION Luciano Floridi
INNOVATION Mark Dodgson and David Gann
INTELLECTUAL PROPERTY Siva Vaidhyanathan
INTELLIGENCE Ian J. Deary
INTERNATIONAL LAW Vaughan Lowe
INTERNATIONAL MIGRATION Khalid Koser
INTERNATIONAL RELATIONS Christian Reus-Smit
INTERNATIONAL SECURITY Christopher S. Browning
INSECTS Simon Leather
IRAN Ali M. Ansari
ISLAM Malise Ruthven
ISLAMIC HISTORY Adam Silverstein
ISLAMIC LAW Mashood A. Baderin
ISOTOPES Rob Ellam
ITALIAN LITERATURE Peter Hainsworth and David Robey
HENRY JAMES Susan L. Mizruchi
JESUS Richard Bauckham
JEWISH HISTORY David N. Myers
JEWISH LITERATURE Ilan Stavans
JOURNALISM Ian Hargreaves
JAMES JOYCE Colin MacCabe
JUDAISM Norman Solomon
JUNG Anthony Stevens
KABBALAH Joseph Dan
KAFKA Ritchie Robertson
KANT Roger Scruton
KEYNES Robert Skidelsky
KIERKEGAARD Patrick Gardiner
KNOWLEDGE Jennifer Nagel
THE KORAN Michael Cook
KOREA Michael J. Seth
LAKES Warwick F. Vincent
LANDSCAPE ARCHITECTURE Ian H. Thompson
LANDSCAPES AND GEOMORPHOLOGY Andrew Goudie and Heather Viles
LANGUAGES Stephen R. Anderson
LATE ANTIQUITY Gillian Clark
LAW Raymond Wacks
THE LAWS OF THERMODYNAMICS Peter Atkins
LEADERSHIP Keith Grint
LEARNING Mark Haselgrove
LEIBNIZ Maria Rosa Antognazza
C. S. LEWIS James Como
LIBERALISM Michael Freeden
LIGHT Ian Walmsley
LINCOLN Allen C. Guelzo
LINGUISTICS Peter Matthews
LITERARY THEORY Jonathan Culler
LOCKE John Dunn
LOGIC Graham Priest
LOVE Ronald de Sousa
MARTIN LUTHER Scott H. Hendrix
MACHIAVELLI Quentin Skinner
MADNESS Andrew Scull
MAGIC Owen Davies
MAGNA CARTA Nicholas Vincent
MAGNETISM Stephen Blundell
MALTHUS Donald Winch
MAMMALS T. S. Kemp
MANAGEMENT John Hendry
NELSON MANDELA Elleke Boehmer
MAO Delia Davin
MARINE BIOLOGY Philip V. Mladenov
MARKETING Kenneth Le Meunier-FitzHugh
THE MARQUIS DE SADE John Phillips
MARTYRDOM Jolyon Mitchell
MARX Peter Singer
MATERIALS Christopher Hall
MATHEMATICAL FINANCE Mark H. A. Davis
MATHEMATICS Timothy Gowers
MATTER Geoff Cottrell
THE MAYA Matthew Restall and Amara Solari
THE MEANING OF LIFE Terry Eagleton
MEASUREMENT David Hand
MEDICAL ETHICS Michael Dunn and Tony Hope
MEDICAL LAW Charles Foster
MEDIEVAL BRITAIN John Gillingham and Ralph A. Griffiths
MEDIEVAL LITERATURE Elaine Treharne
MEDIEVAL PHILOSOPHY John Marenbon
MEMORY Jonathan K. Foster
METAPHYSICS Stephen Mumford
METHODISM William J. Abraham
THE MEXICAN REVOLUTION Alan Knight
MICROBIOLOGY Nicholas P. Money
MICROECONOMICS Avinash Dixit
MICROSCOPY Terence Allen
THE MIDDLE AGES Miri Rubin
MILITARY JUSTICE Eugene R. Fidell
MILITARY STRATEGY Antulio J. Echevarria II
JOHN STUART MILL Gregory Claeys
MINERALS David Vaughan
MIRACLES Yujin Nagasawa
MODERN ARCHITECTURE Adam Sharr
MODERN ART David Cottington
MODERN BRAZIL Anthony W. Pereira
MODERN CHINA Rana Mitter
MODERN DRAMA Kirsten E. Shepherd-Barr
MODERN FRANCE Vanessa R. Schwartz
MODERN INDIA Craig Jeffrey
MODERN IRELAND Senia Pašeta
MODERN ITALY Anna Cento Bull
MODERN JAPAN Christopher Goto-Jones
MODERN LATIN AMERICAN LITERATURE Roberto González Echevarría
MODERN WAR Richard English
MODERNISM Christopher Butler
MOLECULAR BIOLOGY Aysha Divan and Janice A. Royds
MOLECULES Philip Ball
MONASTICISM Stephen J. Davis
THE MONGOLS Morris Rossabi
MONTAIGNE William M. Hamlin
MOONS David A. Rothery
MORMONISM Richard Lyman Bushman
MOUNTAINS Martin F. Price
MUHAMMAD Jonathan A. C. Brown
MULTICULTURALISM Ali Rattansi
MULTILINGUALISM John C. Maher
MUSIC Nicholas Cook
MUSIC AND TECHNOLOGY Mark Katz
MYTH Robert A. Segal
NAPOLEON David Bell
THE NAPOLEONIC WARS Mike Rapport
NATIONALISM Steven Grosby
NATIVE AMERICAN LITERATURE Sean Teuton
NAVIGATION Jim Bennett
NAZI GERMANY Jane Caplan
NEGOTIATION Carrie Menkel-Meadow
NEOLIBERALISM Manfred B. Steger and Ravi K. Roy
NETWORKS Guido Caldarelli and Michele Catanzaro
THE NEW TESTAMENT Luke Timothy Johnson
THE NEW TESTAMENT AS LITERATURE Kyle Keefer
NEWTON Robert Iliffe
NIETZSCHE Michael Tanner
NINETEENTH‑CENTURY BRITAIN Christopher Harvie and H. C. G. Matthew
THE NORMAN CONQUEST George Garnett
NORTH AMERICAN INDIANS Theda Perdue and Michael D. Green
NORTHERN IRELAND Marc Mulholland
NOTHING Frank Close
NUCLEAR PHYSICS Frank Close
NUCLEAR POWER Maxwell Irvine
NUCLEAR WEAPONS Joseph M. Siracusa
NUMBER THEORY Robin Wilson
NUMBERS Peter M. Higgins
NUTRITION David A. Bender
OBJECTIVITY Stephen Gaukroger
OCEANS Dorrik Stow
THE OLD TESTAMENT Michael D. Coogan
THE ORCHESTRA D. Kern Holoman
ORGANIC CHEMISTRY Graham Patrick
ORGANIZATIONS Mary Jo Hatch
ORGANIZED CRIME Georgios A. Antonopoulos and Georgios Papanicolaou
ORTHODOX CHRISTIANITY A. Edward Siecienski
OVID Llewelyn Morgan
PAGANISM Owen Davies
PAKISTAN Pippa Virdee
THE PALESTINIAN-ISRAELI CONFLICT Martin Bunton
PANDEMICS Christian W. McMillen
PARTICLE PHYSICS Frank Close
PAUL E. P. Sanders
IVAN PAVLOV Daniel P. Todes
PEACE Oliver P. Richmond
PENTECOSTALISM William K. Kay
PERCEPTION Brian Rogers
THE PERIODIC TABLE Eric R. Scerri
PHILOSOPHICAL METHOD Timothy Williamson
PHILOSOPHY Edward Craig
PHILOSOPHY IN THE ISLAMIC WORLD Peter Adamson
PHILOSOPHY OF BIOLOGY Samir Okasha
PHILOSOPHY OF LAW Raymond Wacks
PHILOSOPHY OF MIND Barbara Gail Montero
PHILOSOPHY OF PHYSICS David Wallace
PHILOSOPHY OF SCIENCE Samir Okasha
PHILOSOPHY OF RELIGION Tim Bayne
PHOTOGRAPHY Steve Edwards
PHYSICAL CHEMISTRY Peter Atkins
PHYSICS Sidney Perkowitz
PILGRIMAGE Ian Reader
PLAGUE Paul Slack
PLANETARY SYSTEMS Raymond T. Pierrehumbert
PLANETS David A. Rothery
PLANTS Timothy Walker
PLATE TECTONICS Peter Molnar
PLATO Julia Annas
POETRY Bernard O’Donoghue
POLITICAL PHILOSOPHY David Miller
POLITICS Kenneth Minogue
POLYGAMY Sarah M. S. Pearsall
POPULISM Cas Mudde and Cristóbal Rovira Kaltwasser
POSTCOLONIALISM Robert Young
POSTMODERNISM Christopher Butler
POSTSTRUCTURALISM Catherine Belsey
POVERTY Philip N. Jefferson
PREHISTORY Chris Gosden
PRESOCRATIC PHILOSOPHY Catherine Osborne
PRIVACY Raymond Wacks
PROBABILITY John Haigh
PROGRESSIVISM Walter Nugent
PROHIBITION W. J. Rorabaugh
PROJECTS Andrew Davies
PROTESTANTISM Mark A. Noll
PSYCHIATRY Tom Burns
PSYCHOANALYSIS Daniel Pick
PSYCHOLOGY Gillian Butler and Freda McManus
PSYCHOLOGY OF MUSIC Elizabeth Hellmuth Margulis
PSYCHOPATHY Essi Viding
PSYCHOTHERAPY Tom Burns and Eva Burns-Lundgren
PUBLIC ADMINISTRATION Stella Z. Theodoulou and Ravi K. Roy
PUBLIC HEALTH Virginia Berridge
PURITANISM Francis J. Bremer
THE QUAKERS Pink Dandelion
QUANTUM THEORY John Polkinghorne
RACISM Ali Rattansi
RADIOACTIVITY Claudio Tuniz
RASTAFARI Ennis B. Edmonds
READING Belinda Jack
THE REAGAN REVOLUTION Gil Troy
REALITY Jan Westerhoff
RECONSTRUCTION Allen C. Guelzo
THE REFORMATION Peter Marshall
REFUGEES Gil Loescher
RELATIVITY Russell Stannard
RELIGION Thomas A. Tweed
RELIGION IN AMERICA Timothy Beal
THE RENAISSANCE Jerry Brotton
RENAISSANCE ART Geraldine A. Johnson
RENEWABLE ENERGY Nick Jelley
REPTILES T.S. Kemp
REVOLUTIONS Jack A. Goldstone
RHETORIC Richard Toye
RISK Baruch Fischhoff and John Kadvany
RITUAL Barry Stephenson
RIVERS Nick Middleton
ROBOTICS Alan Winfield
ROCKS Jan Zalasiewicz
ROMAN BRITAIN Peter Salway
THE ROMAN EMPIRE Christopher Kelly
THE ROMAN REPUBLIC David M. Gwynn
ROMANTICISM Michael Ferber
ROUSSEAU Robert Wokler
RUSSELL A. C. Grayling
THE RUSSIAN ECONOMY Richard Connolly
RUSSIAN HISTORY Geoffrey Hosking
RUSSIAN LITERATURE Catriona Kelly
THE RUSSIAN REVOLUTION S. A. Smith
SAINTS Simon Yarrow
SAMURAI Michael Wert
SAVANNAS Peter A. Furley
SCEPTICISM Duncan Pritchard
SCHIZOPHRENIA Chris Frith and Eve Johnstone
SCHOPENHAUER Christopher Janaway
SCIENCE AND RELIGION Thomas Dixon and Adam R. Shapiro
SCIENCE FICTION David Seed
THE SCIENTIFIC REVOLUTION Lawrence M. Principe
SCOTLAND Rab Houston
SECULARISM Andrew Copson
SEXUAL SELECTION Marlene Zuk and Leigh W. Simmons
SEXUALITY Véronique Mottier
WILLIAM SHAKESPEARE Stanley Wells
SHAKESPEARE’S COMEDIES Bart van Es
SHAKESPEARE’S SONNETS AND POEMS Jonathan F. S. Post
SHAKESPEARE’S TRAGEDIES Stanley Wells
GEORGE BERNARD SHAW Christopher Wixson
MARY SHELLEY Charlotte Gordon
THE SHORT STORY Andrew Kahn
SIKHISM Eleanor Nesbitt
SILENT FILM Donna Kornhaber
THE SILK ROAD James A. Millward
SLANG Jonathon Green
SLEEP Steven W. Lockley and Russell G. Foster
SMELL Matthew Cobb
ADAM SMITH Christopher J. Berry
SOCIAL AND CULTURAL ANTHROPOLOGY John Monaghan and Peter Just
SOCIAL PSYCHOLOGY Richard J. Crisp
SOCIAL WORK Sally Holland and Jonathan Scourfield
SOCIALISM Michael Newman
SOCIOLINGUISTICS John Edwards
SOCIOLOGY Steve Bruce
SOCRATES C. C. W. Taylor
SOFT MATTER Tom McLeish
SOUND Mike Goldsmith
SOUTHEAST ASIA James R. Rush
THE SOVIET UNION Stephen Lovell
THE SPANISH CIVIL WAR Helen Graham
SPANISH LITERATURE Jo Labanyi
THE SPARTANS Andrew Bayliss
SPINOZA Roger Scruton
SPIRITUALITY Philip Sheldrake
SPORT Mike Cronin
STARS Andrew King
STATISTICS David J. Hand
STEM CELLS Jonathan Slack
STOICISM Brad Inwood
STRUCTURAL ENGINEERING David Blockley
STUART BRITAIN John Morrill
THE SUN Philip Judge
SUPERCONDUCTIVITY Stephen Blundell
SUPERSTITION Stuart Vyse
SYMMETRY Ian Stewart
SYNAESTHESIA Julia Simner
SYNTHETIC BIOLOGY Jamie A. Davies
SYSTEMS BIOLOGY Eberhard O. Voit
TAXATION Stephen Smith
TEETH Peter S. Ungar
TELESCOPES Geoff Cottrell
TERRORISM Charles Townshend
THEATRE Marvin Carlson
THEOLOGY David F. Ford
THINKING AND REASONING Jonathan St B. T. Evans
THOUGHT Tim Bayne
TIBETAN BUDDHISM Matthew T. Kapstein
TIDES David George Bowers and Emyr Martyn Roberts
TIME Jenann Ismael
TOCQUEVILLE Harvey C. Mansfield
LEO TOLSTOY Liza Knapp
TOPOLOGY Richard Earl
TRAGEDY Adrian Poole
TRANSLATION Matthew Reynolds
THE TREATY OF VERSAILLES Michael S. Neiberg
TRIGONOMETRY Glen Van Brummelen
THE TROJAN WAR Eric H. Cline
TRUST Katherine Hawley
THE TUDORS John Guy
TWENTIETH‑CENTURY BRITAIN Kenneth O. Morgan
TYPOGRAPHY Paul Luna
THE UNITED NATIONS Jussi M. Hanhimäki
UNIVERSITIES AND COLLEGES David Palfreyman and Paul Temple
THE U.S. CIVIL WAR Louis P. Masur
THE U.S. CONGRESS Donald A. Ritchie
THE U.S. CONSTITUTION David J. Bodenhamer
THE U.S. SUPREME COURT Linda Greenhouse
UTILITARIANISM Katarzyna de Lazari-Radek and Peter Singer
UTOPIANISM Lyman Tower Sargent
VETERINARY SCIENCE James Yeates
THE VIKINGS Julian D. Richards
VIOLENCE Philip Dwyer
THE VIRGIN MARY Mary Joan Winn Leith
THE VIRTUES Craig A. Boyd and Kevin Timpe
VIRUSES Dorothy H. Crawford
VOLCANOES Michael J. Branney and Jan Zalasiewicz
VOLTAIRE Nicholas Cronk
WAR AND RELIGION Jolyon Mitchell and Joshua Rey
WAR AND TECHNOLOGY Alex Roland
WATER John Finney
WAVES Mike Goldsmith
WEATHER Storm Dunlop
THE WELFARE STATE David Garland
WITCHCRAFT Malcolm Gaskill
WITTGENSTEIN A. C. Grayling
WORK Stephen Fineman
WORLD MUSIC Philip Bohlman
WORLD MYTHOLOGY David Leeming
THE WORLD TRADE ORGANIZATION Amrita Narlikar
WORLD WAR II Gerhard L. Weinberg
WRITING AND SCRIPT Andrew Robinson
ZIONISM Michael Stanislawski
ÉMILE ZOLA Brian Nelson

Available soon:
HANNAH ARENDT Dana Villa
MICROBIOMES Angela E. Douglas
GÖDEL’S THEOREM A. W. Moore
ANSELM Thomas Williams

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Dorothy H. Crawford

VIRUSES
A Very Short Introduction
THIRD EDITION
Great Clarendon Street, Oxford, OX2 6DP, United Kingdom
Oxford University Press is a department of the University of Oxford. It furthers the
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certain other countries
© Dorothy H. Crawford 2011, 2018, 2022
The moral rights of the author have been asserted
First edition published 2011
Second edition published 2018
Third edition published 2022
Impression: 1
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Links to third party websites are provided by Oxford in good faith and for information only.
Oxford disclaims any responsibility for the materials contained in any third party website
referenced in this work.
Contents

Acknowledgments

List of illustrations

List of tables

1 What are viruses?

2 Kill or be killed

3 Emerging virus infections: vertebrate-transmitted viruses

4 Emerging virus infections: arthropod-transmitted viruses

5 Emerging virus infections: the coronaviruses

6 Virus acquisition and eradication

7 Persistent viruses

8 Tumour viruses

9 Viruses past and future

Glossary
Further reading

Publisher’s acknowledgements

Index
Acknowledgements

My sincere thanks go to Dr Karen McAulay for her critical review of


the manuscript and for providing essential information on arthropod-
transmitted viruses.
List of illustrations

1 The structure of viruses


This article was published in Medical Microbiology, 16th edn, D. Greenwood et al.
(eds), p. 23, fig. 2.16 © Elsevier

2 The comparative sizes of a typical bacterium and representative viruses


From L. Collier and J. S. Oxford, Human Virology (OUP, 1993), p. 4, fig. 1.1 © Oxford
University Press

3 The structure of DNA


From B. and D. Charlesworth, Evolution: A Very Short Introduction (OUP, 2003), p. 25,
fig. 5b

4 The retrovirus infectious cycle


© 2017 Takara Bio USA., Inc

5 Molecular evolution of a viral gene over time


Kindly provided by Professor Paul Sharp, University of Edinburgh

6 Portals of virus entry into the human body

7 The Cow-Pock-or-The Wonderful Effects of the New Inoculation, by James


Gillray, 1802
British Cartoon Prints Collection. Prints & Photographs Division, Library of Congress,
LC-USZC4-3147

8 Graphs showing Ebola cases in Guinea, Liberia, and Sierra Leone, 2014–16
Reprinted from Ebola: Mapping the outbreak—BBC News with permission, based on
data taken from WHO Ebola situation reports, data summary, all data and statistics
and Ebola maps Ebola virus disease (who.int)
9 The yellow fever transmission cycle
Source from Fig 15 from D. H. Crawford, The Invisible Enemy (OUP, 2021) © Oxford
University Press

10 Worldwide distribution of Zika virus, dengue fever, and Chikungunya viruses


© NaTHNaC; used with permission of NaTHNaC under Open Government License
v3.0, UK; courtesy of CDC; © 2021, F. Hucke, M. Bestehorn-Willmann & J.J. Bugert,
under exclusive licence to Springer Science Business Media

11 The emergence of SARS in Hong Kong


Used with permission of the Sars Expert Committee

12 Chart showing age-related deaths from COVID-19


From D. H. Crawford, The Invisible Enemy (OUP, 2000), © Oxford University Press

13 CD4 count and viral load during the acute, asymptomatic, and symptomatic
phases of HIV infection
Reproduced from A. Mindel and M. Tenant-Flowers, ‘Natural History and Management
of early HIV infection’, ABC of Aids (2001), with permission from BMJ Publishing Group
Ltd

14 World map showing areas of high incidence of virus-associated tumours


From D. H. Crawford, The Invisible Enemy (OUP, 2000), p. 92, © Oxford University
Press

15 Chart showing the age-related incidence of Burkitt’s lymphoma in Africa


From Hämmerl, L., Colombet, M., Rochford, R. et al. The burden of Burkitt lymphoma
in Africa. Infect Agents Cancer 14, 17 (2019). https://doi.org/10.1186/s13027-019-
0236-7
List of tables

1 Comparison of reproductive number (R0) and herd immunity threshold (HIT)

2 Comparison of SARS-CoV, MERS-CoV, and SARS-CoV-2

3 Cervical cancer incidence rates worldwide in 2018, by region


Data from Statista 2022, Cervical cancer incidence rates by region worldwide 2018 |
Statista
Chapter 1
What are viruses?

The microbe is so very small


You cannot make him out at all,
But many sanguine people hope
To see him through a microscope.
His jointed tongue that lies beneath
A hundred curious rows of teeth;
His seven tufted tails with lots
Of lovely pink and purple spots,
On each of which a pattern stands,
Composed of forty separate bands;
His eyebrows of a tender green;
All these have never yet been seen—
But Scientists, who ought to know,
Assure us that they must be so…
Oh! let us never, never doubt
What nobody is sure about.
‘The Microbe’ (1896), Hilaire Belloc

Primitive microbes evolved on Earth approximately three billion years


ago but were isolated by humans only in the late 19th century,
around 20 years before Hilaire Belloc wrote ‘The Microbe’.

Written to amuse, the poem nonetheless reflects the scepticism of


the times. It must have taken a huge leap of faith for people to
accept that tiny, living organisms were responsible for diseases that
had hitherto been attributed variously to the will of the gods, the
alignment of planets, or miasmic vapours emanating from swamps
and decomposing organic material. Of course, this realization did not
dawn overnight, but as more and more microbes were identified, the
‘germ theory’ took hold, and by the beginning of the 20th century it
was widely accepted even in non-scientific circles that microbes
could cause disease.

Key to this momentous leap in understanding were technical


developments in microscopes made by the Dutch lens-maker Antonie
van Leeuwenhoek (1632–1723) in the 17th century. He was the first
to visualize microbes, but it was not until the mid-1800s that Louis
Pasteur (1822–95) working in Paris and Robert Koch (1843–1910) in
Berlin established ‘germs’ as the cause of infectious diseases.

Koch discovered the first bacterium, Bacillus anthraci, in 1876, and


soon the causative microbes of feared diseases like anthrax,
tuberculosis, cholera, diphtheria, tetanus, and syphilis were
identified and characterized. But these were all caused by bacteria,
and it soon became clear that there remained a group of infectious
diseases for which causative organisms could not be found, such as
smallpox, measles, mumps, rubella, and flu. These microbes were
obviously very small as they passed through filters that trapped
bacteria, and in consequence were called ‘filterable agents’. At the
time, most scientists thought these were just tiny bacteria. It was
not until the invention of the electron microscope in 1939, able to
magnify over 100,000 times, that viruses were first visualized and
their structure elucidated, showing them to be a unique class of
microbes.

Viruses are not cells but particles. They consist of a protein coat, the
capsid, surrounding and protecting a piece of genetic material. The
whole structure is called a virion. Capsids come in various shapes
and sizes, each characteristic of the family to which a virus belongs.
They are built up of protein subunits called capsomeres and it is the
arrangement of these around the central genetic material that
determines the shape of the virion. For example, pox viruses are
brick-shaped, herpes viruses are icosahedral (20-sided polygons),
the rabies virus is bullet-shaped, and the tobacco mosaic virus is
long and thin like a rod (Figure 1).
1. The structure of viruses.

Some viruses have an outer layer surrounding the capsid called an


envelope, which they acquire when budding through the cell surface
membrane.

Most viruses are too small to be seen under a light microscope. In


general, they vary in size from 20 to 300 nanometres (nm) in
diameter (1 nm is a thousand millionth of a metre) (Figure 2).
2. The comparative sizes of a typical bacterium and representative
viruses.

However, the genus of so-called giant viruses forms an exception.


Mimivirus (short for ‘microbe-mimicking virus’) was the first of these
to be discovered, in 1992, and measures 600 nm; larger than some
bacteria.

Inside the virus capsid is the genetic material, or genome, which is


either RNA or DNA depending on the type of virus (Figure 3). The
genome contains the virus’s genes, which carry the code for making
new viruses, and transmits their inherited characteristics to the next
generation. Viruses usually have between 4 and 200 genes, but
again the giant viruses are unusual in having an estimated 600 to
1,000 genes, more than many bacteria.
3. The structure of DNA, showing the two complementary strands that
form the helix. The backbone of each strand is composed of molecules
of the sugar deoxyribose (S) that are linked to each other through
phosphate molecules (P). Each sugar is connected to a nucleotide
molecule, and these form the ‘letters’ of the genetic alphabet. These
are: adenine (A), guanine (G), cytosine (C), and thymine (T). The
structure of RNA is similar to DNA but its nucleotides are adenine,
guanine, cytosine, and uracil.

Once it was appreciated that viruses carried either DNA or RNA, but
never both, a system of classification was devised based on the
following criteria to assign viruses into families, genera, and species:

• the type of nucleic acid (DNA or RNA);


• the shape of the virus capsid;
• the capsid diameter and/or number of capsomeres;
• the presence or absence of a viral envelope.

Since the early 1980s, when the first virus genome was fully
mapped, genome sequencing has become a routine technique that
provides valuable information for virus classification. Indeed, with
increasingly sophisticated methods for virus discovery, many viruses
are now identified long before their actual physical structure is
visualized. In these cases, the molecular structure of the DNA or
RNA is compared with that of other known viruses to assign the new
virus to a family.

Cells of free-living organisms, including bacteria, contain a variety of


organelles essential for life such as ribosomes that manufacture
proteins, mitochondria, or other structures that generate energy, and
complex membranes for transporting molecules within the cell across
the cell wall. Viruses, not being cells, have none of these, and are
therefore inert until they infect a living cell. Once inside, they hijack
the cell’s organelles and use what they need to complete the virus
life cycle, often killing the cell in the process. As such they are called
obligate parasites. Even mimivirus, which infects amoebae, must
borrow the amoeba’s organelles to manufacture its proteins in order
to assemble new mimiviruses.

Plant viruses either enter cells through a break in the cell wall or are
injected by sap-sucking insect vectors like aphids. They then spread
very efficiently from cell to cell via plasmodesmata, pores that
transport molecules between cells. In contrast, animal viruses infect
cells by binding to specific cell surface receptor molecules. The cell
receptor is like a lock, and only viruses that carry the right receptor-
binding key can open it and enter that particular cell. Receptor
molecules differ from one type of virus to another, and while some
are found on many cell types, others have a much more restricted
distribution. This is exemplified by the two pandemic viruses: human
immunodeficiency virus (HIV), the cause of acquired
immunodeficiency syndrome (AIDS), and severe acute respiratory
syndrome-coronavirus-2 (SARS-CoV-2), the cause of COVID-19. The
cell receptor for HIV is the CD4 molecule, which is mainly expressed
on immune cells called helper T cells. This specific interaction
defines the outcome of HIV infection, since it leads to destruction of
CD4-positive ‘T’ cells that are critical to the immune response.
Without antiviral treatment the immune system eventually fails, and
lethal opportunistic infections ensue. In contrast, the cell receptor for
SARS-CoV-2, ACE-2 (angiotensin-converting enzyme-2), is widely
expressed in the body, particularly on cells in the respiratory tract,
blood vessels, heart, and kidneys, and this distribution determines
the manifestations of severe COVID-19 (see Chapter 5).

Once a virus has bound to its cellular receptor, the capsid penetrates
the cell and its genome (DNA or RNA) is released into the cell
cytoplasm. The main ‘aim’ of a virus is to reproduce successfully, and
to do this its genetic material must download the information it
carries. Mostly, this will take place in the cell’s nucleus, where the
virus can access the molecules it requires to begin manufacturing its
own proteins. Some large viruses, like pox viruses, carry genes for
the enzymes they need to make their proteins and so are more self-
sufficient and can complete the whole life cycle in the cytoplasm.

Inside a cell DNA viruses simply masquerade as pieces of cellular


DNA, and their genes are transcribed and translated using as much
of the cell’s machinery as they require. The viral DNA code is
transcribed into RNA messages (transcription) which are read and
translated into individual viral proteins (translation) by the cell’s
ribosomes.

The separate virus components are then assembled into thousands


of new viruses which are often so tightly packed inside the cell that
it eventually bursts open and releases them, inevitably killing the
cell. Alternatively, new viruses leave rather more sedately by
budding through the cell membrane and in doing so acquire an
envelope. In the latter case, the cell may survive and act as a
reservoir of infection.

RNA viruses are one step ahead of DNA viruses in already having
their genetic code as RNA. As they carry enzymes that enable their
RNA to be copied and translated into proteins, they are not so
dependent on cellular enzymes and can often complete their life
cycle in the cytoplasm without causing major disruption to the cell.

Retroviruses are a family of RNA viruses, including HIV, that have


evolved a unique trick for establishing a lifelong infection of a cell
while hiding from immune attack. Retrovirus particles contain an
enzyme called reverse transcriptase which, once inside a cell,
converts their RNA to DNA (Figure 4). This viral DNA can then join
into the cell’s DNA chain using another viral enzyme called integrase;
a process called integration. The integrated viral sequence is called
the provirus, and is effectively archived in the cell, remaining there
permanently to be copied along with cellular DNA when the cell
divides. The provirus is inherited by the two daughter cells, so
building up a store of infected cells inside its host. At any time, a
provirus can manufacture new viruses which bud from the cell
surface, but in this instance it kills the cell.

4. The retrovirus infectious cycle, showing viral entry into a cell


followed by reverse transcription, integration, transcription, and
translation of the genome, virus assembly, and budding of new particles
from the cell surface.

In mammalian cells, the process of copying DNA during cell division


is highly regulated, with a proof-reading system and checkpoints in
place to detect damaged or miscopied DNA and to correct the
mistakes. If the damage is too great, cells have an ‘auto-destruct’
programme called apoptosis that induces cell death rather than
allowing the cell to pass on its faulty DNA. Despite these checks,
mistakes slip through, causing mutations to be replicated and passed
on to future generations (Figure 5).
5. Molecular evolution of a viral gene over time. The extent of
accumulated difference between the sequences is used to construct an
evolutionary tree, in which the lengths of the horizontal branches are
drawn to scale, and denote the time since common ancestors (shown as
circles).

Virus genomes mutate far more rapidly than the human genome,
partly because viruses reproduce in a day or two with many
thousands of offspring. Also, RNA viruses generally have no proof-
reading system so they have a higher mutation rate than DNA
viruses.

Thus, every time a virus infects a cell, its DNA or RNA may be copied
thousands of times, and as each new strand is incorporated into a
new virus particle, every round of infection throws up several mutant
viruses. This high mutation rate in viruses is their lifeline; in some, it
is essential for their survival. Each round of infection produces some
viruses that are non-viable due to mutations that interrupt the
function of essential genes, and others with mutations that cause no
change in function. But a few of the offspring will have beneficial
mutations, giving them a selective advantage over their siblings. The
benefit may be due to any number of changes, including a
heightened ability to hide from immune attack; to infect more easily;
to resist antiviral drugs; or to reproduce at a faster rate. Whatever
the advantage, it will lead to that particular mutant virus outstripping
its siblings and eventually taking over in the population. Measles
virus, for example, has been infecting the human population for at
least 2,000 years, but scientists calculate that the present-day
measles strain arose much more recently (see Chapter 7).
Presumably, this virus strain was ‘fitter’ than its predecessor in some
way and so eventually replaced the former strain worldwide. And as
I write we are regularly experiencing the emergence of new strains
of SARS-CoV-2 during the COVID-19 pandemic, with strains better at
transmission between hosts replacing their predecessors globally
(see Chapter 5).

Analysing the mutations in a virus’s genome is a useful way of


tracking its history. The molecular clock hypothesis, which was
developed in the 1960s, states that the mutation rate per generation
is constant for any given gene. In other words, as applied to viruses,
two samples of the same virus isolated at the same time from
different sources will have evolved for the same length of time since
their common ancestor (Figure 5). Because they will both have been
accumulating mutations at a constant rate, the degree of difference
between their gene sequences provides a measure of the time that
has passed since their common ancestor. Scientists use the
molecular clock to estimate the time of origin of certain viruses, and
plot evolutionary (or phylogenetic) trees showing their degree of
relatedness to other viruses. Because viruses have a high mutation
rate, significant evolutionary change, estimated at around 1 per cent
per year for HIV, can be measured over a short timescale. This
technique was instrumental in uncovering the animal origin of HIV
and is at present being used to search for the origin of SARS-CoV-2.
Because virus particles are inert, lacking the ability to generate
energy or manufacture proteins independently, they are not
generally regarded as living organisms. Nonetheless, they are pieces
of genetic material that parasitize cells, very efficiently exploiting the
cells’ internal machinery to reproduce themselves. So how and when
did these cellular hijackers originate?

We do not yet know the answer to this question, but there are three
theories—the virus-first, the progressive, and the regressive
theories, none of which explains the various features of all the
viruses we observe today. It is now generally accepted that viruses
are truly ancient. The fact that viruses sharing common features
infect organisms in all three domains of life—Archaea, Bacteria, and
Eukarya—suggests that they evolved before these domains
separated from their common ancestor, called the ‘last universal
cellular ancestor’ (LUCA). It is also generally accepted that the first
replicating molecule to evolve was RNA rather than DNA. So the
virus-first theory suggests that RNA viruses evolved before more
complex cells although they now depend on them for their survival.
Then DNA viruses may have evolved from their more ancient RNA
counterparts. This suggestion is supported by the existence of
retroviruses, with their ability to transcribe RNA into DNA. In so
doing, they reverse the more usual flow of genetic information from
DNA to RNA to protein.

The regressive theory proposes that large DNA viruses, for example
pox and mimiviruses, may represent previously free-living life forms
that have now lost their ability to reproduce independently. In
contrast the progressive theory suggests that viruses derived from
escaped fragments of genetic material that acquired a protein coat
and became infectious.

Whatever the truth of it, none of these theories can explain the
evolution of both RNA and DNA viruses, which may therefore have
evolved separately.
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of men. It was as though they were back in the days of the old
Hebrew prophets when the hand of the Lord stretched out and laid
itself upon wicked men for their punishment when the measure of
their time was full.
“He tried to stand above the law in this valley,” Hollister told her. “He
wanted to stop progress—said there shouldn’t be any dam to reclaim
the Flat Tops for settlers. Merrick will rebuild it. The land will be
watered. Your ranch will be good as ever in three months. And he’ll
be buried and forgotten.”
“And poor Don Black?” she whispered. “Poor Don, who never had a
chance in this world, or, if he had one, muddled it so badly?”
He could only hope that Don had gone to a better-ordered world
where circumstances did not dominate good intentions.
Betty’s sense of tragedy lingered just now no longer than a cinema
picture. The life urge in her clamored for expression. No world could
be a sad one with her and Tug in it.
“Shall I go in and tell your father now?” the young man asked.
“Soon.” She made a rustling little motion toward him and found
herself in his arms. “Isn’t it splendid, boy? To-day’s the best ever, and
to-morrow will be better than to-day—oh, heaps better—and after
that all the years forever and ever.”
He looked into the deep lustrous eyes of his straight slim girl. What a
wife she would be! How eager and provocative, this white flame of
youth so simple and so complex! Her happiness now would be in his
hands. The responsibility awed him, filled him with a sense of
solemnity.
“Forever is a long time,” he said, smiling, and quoted a stanza of
magazine verse they had lately read together.
It began, “How far will you go with me, my love?” Close-held in his
arms, Betty answered without a moment of hesitation.
“She smiled at the stile with a sweet disdain;
She scoffed at the bridge and the great oak tree;
And looked me full in the eyes and said:
‘I will go to the end of the lane with thee.’”
The door of the inner room opened and Clint stood on the threshold.
“Hello!” he said, surprised.
Betty disengaged herself, blushing. “He’s decided to take me after
all, Dad,” she said demurely.
“Hmp! Has he? Kinda looks that way.” Clint gripped Hollister’s hand
till it hurt. It was the best he could do just now to show the gratitude
he felt for what this man had done.
“That’s not quite the way I put it, sir,” Tug said.
“Doesn’t matter how you put it, boy. It’ll be her say-so from now on.
Don’t I know her? Hasn’t she bossed me scandalous since she was
knee-high to a gosling?”
“Now, Dad, you’re giving me a bad name,” Betty protested, hugging
her father.
“If he ain’t man enough to stand some bossing, he’d better quit right
now before he says, ‘I do.’”
“He likes being bossed, Dad,” Betty announced, and the imps of
deviltry were kicking up their heels in her eyes. “Don’t you, Tug?”
Hollister looked at the girl and smiled. “I’ll say I do,” he admitted.
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for his soul.
WEBSTER: MAN’S MAN
In a little Jim Crow Republic in Central America, a man
and a woman, hailing from the “States,” met up with a
revolution and for a while adventures and excitement
came so thick and fast that their love affair had to wait for
a lull in the game.
CAPTAIN SCRAGGS
This sea yarn recounts the adventures of three rapscallion
sea-faring men—a Captain Scraggs, owner of the green
vegetable freighter Maggie, Gibney the mate and
McGuffney the engineer.
THE LONG CHANCE
A story fresh from the heart of the West, of San Pasqual, a
sun-baked desert town, of Harley P. Hennage, the best
gambler, the best and worst man of San Pasqual and of
lovely Donna.

Grosset & Dunlap, Publishers, New York


JACKSON GREGORY’S NOVELS
May be had wherever books are sold. Ask for Grosset &
Dunlap’s list.

THE EVERLASTING WHISPER


The story of a strong man’s struggle against savage
nature and humanity, and of a beautiful girl’s regeneration
from a spoiled child of wealth into a courageous strong-
willed woman.
DESERT VALLEY
A college professor sets out with his daughter to find gold.
They meet a rancher who loses his heart, and become
involved in a feud. An intensely exciting story.
MAN TO MAN
Encircled with enemies, distrusted, Steve defends his
rights. How he won his game and the girl he loved is the
story filled with breathless situations.
THE BELLS OF SAN JUAN
Dr. Virginia Page is forced to go with the sheriff on a night
journey into the strongholds of a lawless band. Thrills and
excitement sweep the reader along to the end.
JUDITH OF BLUE LAKE RANCH
Judith Sanford part owner of a cattle ranch realizes she is
being robbed by her foreman. How, with the help of Bud
Lee, she checkmates Trevor’s scheme makes fascinating
reading.
THE SHORT CUT
Wayne is suspected of killing his brother after a violent
quarrel. Financial complications, villains, a horse-race and
beautiful Wanda, all go to make up a thrilling romance.
THE JOYOUS TROUBLE MAKER
A reporter sets up housekeeping close to Beatrice’s
Ranch much to her chagrin. There is “another man” who
complicates matters, but all turns out as it should in this
tale of romance and adventure.
SIX FEET FOUR
Beatrice Waverly is robbed of $5,000 and suspicion
fastens upon Buck Thornton, but she soon realizes he is
not guilty. Intensely exciting, here is a real story of the
Great Far West.
WOLF BREED
No Luck Drennan had grown hard through loss of faith in
men he had trusted. A woman hater and sharp of tongue,
he finds a match in Ygerne whose clever fencing wins the
admiration and love of the “Lone Wolf.”

Grosset & Dunlap, Publishers, New York


EMERSON HOUGH’S NOVELS
May be had wherever books are sold. Ask for Grosset
and Dunlap’s list.

THE COVERED WAGON


NORTH OF 36
THE WAY OF A MAN
THE STORY OF THE OUTLAW
THE SAGEBRUSHER
THE GIRL AT THE HALFWAY HOUSE
THE WAY OUT
THE MAN NEXT DOOR
THE MAGNIFICENT ADVENTURE
THE BROKEN GATE
THE STORY OF THE COWBOY
THE WAY TO THE WEST
54-40 OR FIGHT
HEART’S DESIRE
THE MISSISSIPPI BUBBLE
THE PURCHASE PRICE

GROSSET & DUNLAP, Publishers, NEW


YORK
GEORGE W. OGDEN’S WESTERN
NOVELS
May be had wherever books are sold. Ask for Grosset &
Dunlap’s list.

THE BARON OF DIAMOND TAIL


The Elk Mountain Cattle Co. had not paid a dividend in
years; so Edgar Barrett, fresh from the navy, was sent
West to see what was wrong at the ranch. The tale of this
tenderfoot outwitting the buckaroos at their own play will
sweep you into the action of this salient western novel.
THE BONDBOY
Joe Newbolt, bound out by force of family conditions to
work for a number of years, is accused of murder and
circumstances are against him. His mouth is sealed; he
cannot, as a gentleman, utter the words that would clear
him. A dramatic, romantic tale of intense interest.
CLAIM NUMBER ONE
Dr. Warren Slavens drew claim number one, which entitled
him to first choice of rich lands on an Indian reservation in
Wyoming. It meant a fortune; but before he established his
ownership he had a hard battle with crooks and politicians.
THE DUKE OF CHIMNEY BUTTE
When Jerry Lambert, “the Duke,” attempts to safeguard
the cattle ranch of Vesta Philbrook from thieving
neighbors, his work is appallingly handicapped because of
Grace Kerr, one of the chief agitators, and a deadly enemy
of Vesta’s. A stirring tale of brave deeds, gun-play and a
love that shines above all.
THE FLOCKMASTER OF POISON CREEK
John Mackenzie trod the trail from Jasper to the great
sheep country where fortunes were being made by the
flock-masters. Shepherding was not a peaceful pursuit in
those bygone days. Adventure met him at every turn—
there is a girl of course—men fight their best fights for a
woman—it is an epic of the sheeplands.
THE LAND OF LAST CHANCE
Jim Timberlake and Capt. David Scott waited with restless
thousands on the Oklahoma line for the signal to dash
across the border. How the city of Victory arose overnight
on the plains, how people savagely defended their claims
against the “sooners;” how good men and bad played
politics, makes a strong story of growth and American
initiative.
TRAIL’S END
Ascalon was the end of the trail for thirsty cowboys who
gave vent to their pent-up feelings without restraint. Calvin
Morgan was not concerned with its wickedness until Seth
Craddock’s malevolence directed itself against him. He did
not emerge from the maelstrom until he had obliterated
every vestige of lawlessness, and assured himself of the
safety of a certain dark-eyed girl.

Ask for Complete free list of G. & D. Popular Copyrighted Fiction

GROSSET & DUNLAP, Publishers, NEW


YORK
TRANSCRIBER’S NOTES:
Obvious typographical errors have been corrected.
Inconsistencies in hyphenation have been
standardized.
Archaic or variant spelling has been retained.
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