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Genes: 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 d
­ iscipline. 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 AMERICAN IMMIGRATION

THE ABRAHAMIC RELIGIONS
Charles L. Cohen
ACCOUNTING Christopher Nobes
ADDICTION Keith Humphreys
ADOLESCENCE Peter K. Smith
THEODOR W. ADORNO
Andrew Bowie
ADVERTISING Winston Fletcher
AERIAL WARFARE Frank Ledwidge
AESTHETICS Bence Nanay
AFRICAN AMERICAN HISTORY
Jonathan Scott Holloway
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
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
ANCIENT GREEK AND ROMAN
SCIENCE Liba Taub
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
ANSELM Thomas Williams
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
HANNAH ARENDT Dana Villa
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
THE BOHEMIANS David Weir
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
BRITISH CINEMA Charles Barr
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
THE CIVIL RIGHTS MOVEMENT
Thomas C. Holt
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
CONDENSED MATTER PHYSICS
Ross H. McKenzie
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 Naomi Zack
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 G. Stackhouse Jr.
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
GÖDEL’S THEOREM A. W. Moore
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 L. 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 EMOTIONS
Thomas Dixon
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
JAPANESE LITERATURE Alan Tansman
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
THE JURY Renée Lettow Lerner
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 ANALYSIS
Richard Earl
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
MICROBIOMES Angela E. Douglas
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
NANOTECHNOLOGY Philip Moriarty
NAPOLEON David A. 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
OBSERVATIONAL ASTRONOMY
Geoff Cottrell
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 J. C. 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
PSEUDOSCIENCE Michael D. Gordin.
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
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 Jonathan Slack

GENES
A Very Short Introduction
second edition
 Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
Oxford University Press is a department of the University of Oxford.
It furthers the University’s objective of excellence in research, scholarship,
and education by publishing worldwide. Oxford is a registered trade mark of
Oxford University Press in the UK and in certain other countries
© Jonathan Slack 2023
The moral rights of the author have been asserted
First edition published 2014
This edition published 2023
All rights reserved. No part of this publication may be reproduced, stored in
a retrieval system, or transmitted, in any form or by any means, without the
prior permission in writing of Oxford University Press, or as expressly permitted
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and you must impose this same condition on any acquirer
Published in the United States of America by Oxford University Press
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Library of Congress Control Number: 2023930635
ISBN 978–0–19–285670–8
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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.
 Comments on the first edition

‘This is a concise and accurate account of genes, and what they

are, in a readable and convenient format.’
Sir Paul Nurse, Director, Francis Crick Institute

‘Unlike the genetic material itself, this book is short, sharp, and
to the point.’
Steve Jones, University College London

‘We all need to know what genes are. Slack tells us with
authority, clarity and grace.’
Armand Leroi, Imperial College, London

‘The essential guide for getting up to speed with the

­ ver-­changing and crazily complex science of genetics.’
e
Adam Rutherford, University College London

‘With more heat than light in many areas surrounding genetics,

it is refreshing to have such a concise, precise and matter-­of-­fact
introduction to the field. This is strongly recommended to
anyone with even a passing interest in a field set to dominate
our lives.’
Laurence Hurst, Director of The Milner Centre for Evolution,
University of Bath
Contents

Preface xix

List of illustrations xxi

1 Genes before 1944 1

2 Genes as DNA 17

3 Mutations and gene variants 40

4 Genes as markers 60

5 Genes of small effect 87

6 Genes in evolution 109

Conclusion: the varied concepts of the gene 129

Glossary 133

Further reading 139

Index 143
Preface

What is a gene? In essence it is a molecule of DNA, present in

every one of our cells, that controls the synthesis of one particular
protein in our bodies. But this simple definition does not do
justice to the richness of the gene concept and its many
ramifications throughout the life sciences. For example, some
genes do not code for proteins, some appear to have no function at
all, others are presumed to exist but have not been identified in
terms of DNA.

Everyone has heard of genes and we have come to feel that they
are fundamental to who we are. But there is also a lot of
uncertainty and confusion about what we are really referring to
and what it means in a particular context. For example, does
having cancer in a first degree relative mean I have ‘cancer genes’?
If I have genes also found in Stone Age fossils, does this mean
I am very primitive? Does the existence of ‘selfish genes’ mean that
human nature is inherently selfish?

We also know that genes have become the basis of a huge

technology which generates pharmaceuticals, diagnostic tests,
forensic identification, paternity tests, and GM crops, which are
welcomed by some people and feared by others. Many are
suspicious of the gene concept as applied to ethnicity, intelligence,
criminality, or other human attributes, while others presume that
 most variation of these things is due to genetic differences
between people. Despite their preconceptions, many have a desire
to understand this entity which penetrates so deeply into their
lives but still seems rather mysterious and confusing.

This book is not a textbook of genetics, but it is a brief

introduction to the various conceptions of the gene currently
used in the life sciences. These conceptions are quite distinct from
each other. The ‘gene’ of molecular biology is, indeed, a piece of
DNA that encodes a protein, or sometimes a functional RNA
(ribonucleic acid). The ‘genes’ responsible for attributes such as
human height or the incidence of many diseases are differences
between DNA sequences that make up the same coding sequence
in different individuals. The ‘genes’ in studies of ancestry are other
differences in DNA sequence that are mostly not in coding genes
at all. The ‘genes’ of evolutionary biology or sociobiology are
hypothetical entities imagined to provide the basis for theories
of how certain traits can arise in evolution. Most of these have
Genes

not been identified in terms of DNA.

In what follows, some flesh will be put on each of these different

concepts of the gene. The aim is to enable readers to appreciate
the main ideas about genes, to evaluate contentious issues, and to
navigate to more advanced texts if they wish. Terms introduced
in italics are explained in the Glossary at the end.

I am indebted to the following for reading drafts and commenting

on the manuscript: Helen Brittain, Jill Maidment, Janet Slack,
and Zoe Yeomans.

xx
List of illustrations
1 New Leicester sheep. From
David Low’s The Breeds of
Domestic Animals of the British
Islands, London, 1842 3
Heritage Images/Glowimages.com
2 Mendel’s peas 6
3 DNA two-­dimensional
structure 13
4 DNA three-­dimensional
structure 19
Creative Commons
Attribution-Share Alike 3.0
5 Protein synthesis 21
6 Gene structure 24
7 Behaviour of chromosomes
on cell division 26
From Jonathan M. W. Slack,
Essential Developmental Biology,
3rd Edition, John Wiley & Sons
8 Development 30
From Jonathan M. W. Slack,
Essential Developmental Biology,
3rd Edition, John Wiley & Sons
9 The effect of gene drive
on a population 39
Springer Nature
10 Keto-enol 42
11 Romanov family 52
Heritage Images/Glowimages.com
12 Common ancestors 73
13 Human migrations out of
Africa as revealed by the
study of genetic markers
in populations 75
14 A Neanderthal face
reconstructed by anatomical
methods and a Denisovan face
reconstructed from the skull
structure deduced from the
DNA methylation pattern 85
Maayan Harel (14b)
 15 Tall and short 95 18 Altruistic behaviour by the
AFP/Getty Images female nighthawk 121

16 Genetic drift 111 19 Hypothetical genetics

of altruism in social
17 Tree of life 113 insects 123
Genes

xxii
Chapter 1
Genes before 1944

In 1938 a remarkable pair of articles was published in the

Quarterly Review of Biology. They were by a little-­known
American biology professor from the University of Missouri,
Addison Gulick, and they were about the nature of the gene. These
articles are rarely consulted today since they were written shortly
before it was discovered that genes consisted of deoxyribonucleic
acid (DNA). However, they are remarkable in showing how much
was known about genes even before their chemical nature was
established. Gulick knew that genes were located in the
chromosomes of the cell nucleus, and were complex structures that
somehow directed the synthesis of enzymes and the development
of the organism. He knew that they normally remained stable
from one generation to the next, and that occasional changes,
called mutations, could spread through the population and be the
basis of evolution by natural selection. He also made surprisingly
accurate estimates of the sizes and numbers of genes in various
types of organism. These articles illustrate that to appreciate how
our current understanding of the gene came about we need to go
back much further than the famous ‘double helix’ structure of
DNA, discovered in 1953.

Two completely separate lines of work led to our modern view,

and they came together shortly after the appearance of Addison
Gulick’s articles to create the new science of molecular biology.
1
 One was the study of heredity by biological experimentation and
the other was the study of the chemistry of DNA.

Biology of heredity
Before the 18th century there was little informed speculation
about heredity. Even the word did not exist (‘heredité’ first
appeared in France, ‘genetic’ in England, both about 1830).
Historically there had been plenty of animal breeding and vague
ideas of ‘blood lines’, but this was uninformed by much
understanding of reproduction. Although selective breeding in
agriculture had been happening for thousands of years, the first
systematic applications took place in the 18th century. Robert
Bakewell, a sheep breeder from Dishley, near Loughborough,
England, bred a line of New Leicester sheep that grew faster
and produced more meat than before (Figure 1). This was done
by mating the best yielding males and females to create a
self-­reproducing population (breed) that maintained the new
Genes

characteristics in a stable way. The experience of animal breeding

conveyed the idea that heredity involved the blending, or
averaging, of the distinct characteristics, also known as traits, of
the parents. It was quite understandable that animal breeders
should have believed in the blending of traits since this is what
you see when animals are mated and characters such as height or
weight or growth rate are measured. But the blending theory of
inheritance was to become a serious problem for Darwin’s theory
of natural selection.

By the time of Darwin’s work in the mid-­19th century, the fact that
biological evolution had occurred was already accepted by some
scientists, mostly on the basis of the changes of the types of animal
or plant remains seen at different levels of the fossil record. The
real impact of Darwin’s work, and that of his contemporary Alfred
Wallace, was to provide an actual and credible mechanism for the
changes seen in living organisms over evolutionary time. This
mechanism was natural selection, and the case for it is simply
2
 Genes before 1944
1. New Leicester sheep. From David Low’s The Breeds of Domestic
Animals of the British Islands, London, 1842.

stated. If a population of animals or plants varies with regard to

some traits, if those traits are heritable, and if they affect the
likelihood of reproduction, then the composition of the population
will inevitably shift between each generation.

The traits associated with more reproduction will become more

common, and will eventually displace the alternatives. The
direction and speed of the shift will be determined by the selective
conditions that cause the differential reproduction of the
individuals with the different traits. The theory of natural
selection seems very compelling, and it is especially compelling as
presented by Darwin in The Origin of Species (1859), which
contains a huge array of examples drawn from natural history to
support the case. At the time, the main opposition to the theory
was from religious groups who realized that the principle of
3
 natural selection undermined the ‘argument from design’, an
important argument for the existence of God, and also from those
offended by the idea of a biological kinship between humans and
animals. However, there was also some scientific opposition; the
most serious was that which focused on the difficulty of
reconciling natural selection with blending inheritance.

Supposing that one individual is slightly better suited to

reproduction than others due to the possession of a particular trait
(e.g. they might be able to digest some additional component in
their food), he or she will most likely mate with an individual
lacking it and their offspring will then have it in a diluted form, as
the favourable trait is rare. After three or four generations the
hereditary factors responsible for the trait will be diluted out
almost completely. Therefore selection has only a few generations
to operate and this will not be enough to change the whole
breeding population unless the reproductive advantage conferred
by the new trait is very large indeed. Darwin himself was well
Genes

aware of the problem but he was also opposed to the idea of large
jumps in evolution and favoured the idea that evolution
progressed in a smooth and imperceptible manner via many
small changes.

Some thinkers followed this argument to its logical conclusion

and concluded that the hereditary factors responsible for
evolution must have large effects, such that substantial selection
could occur before they became diluted out. In this way the trait
might become common enough for some matings to occur
between parents who both possessed it and it would no longer be
diluted in their offspring. Among these thinkers was William
Bateson, who collected in his book Materials for the Study of
Variation (1894) a remarkable set of examples of discontinuous
and qualitative variation within animal and plant populations.
More direct evidence for the existence of large heritable changes
came from observation of spontaneously occurring ‘mutations’. In
particular the Dutch botanist Hugo de Vries in 1886 observed the
4
de novo appearance of dramatically new forms of the Evening
Primrose, which bred true in subsequent generations. Despite
these findings, blending inheritance remained a serious problem
for the theory of natural selection.

In fact the solution to the problem had been provided as early as

1866 by Gregor Mendel, a monk at the Abbey of St Thomas in
Brno, now in the Czech Republic. In the early 19th century Brno
was a centre of textile manufacture and of sheep breeding, and the
Abbey already had a two-­hectare experimental garden. Mendel
had received education about animal and plant breeding in the
course of his studies of philosophy at the University of Olomouc,
and he was encouraged to continue his work at Brno by the Abbot.
Between 1856 and 1863, Mendel conducted a number of
experiments with peas. He was fortunate enough to choose simple
traits, which we would now call traits determined by single genes,

Genes before 1944

rather than complex traits determined by many genes. Among
these traits were a round or wrinkled appearance, and a green or
yellow colour. Mendel postulated that there were invisible
hereditary ‘factors’ causing each visible trait, and showed that
there were predictable rules for their inheritance. His breeding
experiments indicated that each individual plant contained two
factors for each trait, one derived from each parent. When
reproductive cells (pollen or eggs) are formed, each contains just
one factor, randomly selected from the two possibilities available
in that plant. In some cases one factor would suppress the other:
we should now call this a dominant gene. For example, a cross
between yellow and green peas gives only yellow offspring.
However, if these offspring are crossed to each other, then
25 per cent of the next generation are green, indicating that the
green factors are still there, but cannot be expressed in the
presence of the yellow factors (Figure 2). The green factor would
now be called a recessive gene. So, Mendel showed that the
hereditary factors behaved as discrete units, such that each parent
provided one to each offspring and the appearance of the offspring
depended on the specific combination of factors inherited and the
5
 Yellow (YY) Yellow (Yg)

(Yg) (Yg) (YY) (Yg)

Green (gg) Yellow (Yg)

(Yg) (Yg) (Yg) (gg)

First generation Second generation

2. Mendel’s peas. When yellow peas are crossed to green peas, the first
generation seeds are all yellow. But when members of the first
generation are crossed to each other, 25 per cent of the second
generation seeds are green. Mendel explained this by postulating
hereditary factors, here called Y for yellow and g for green, such that
the yellow factor is dominant over the green where they occur together.

dominance rules between them. Mendel published his work in the

Verhandlungen des Naturforschenden Vereines in Brünn in 1866.
Genes

But this was what we should now call a ‘low-­impact journal’, and
nobody noticed. After he became Abbot in 1868 he was mostly
occupied with administrative duties, and he died in 1884 with the
wider world still ignorant of the founding principles of genetics
that he had discovered.

The 20th century

In Western Europe and America, arguments continued over
whether natural selection could work through blending
inheritance, and whether mutations of large effect were a
credible source of variation. Not until 1900 was Mendel’s work
rediscovered and further developed. Several people were
responsible for this including Hugo de Vries of mutation fame and
the German botanist Carl Correns. It was rapidly realized that the
major contradiction of blending inheritance had now been
removed. Instead of being diluted out in each generation,
Mendel’s factors were stable and persisted from generation to
6
generation. The variation in a population existed because of the
differences between the hereditary factors that were present in
each individual. Recessive traits could appear in offspring due to
the random assortment of factors at reproduction, so it was not
necessary to postulate new mutations to explain every newly
appearing variation. Moreover the more complex characteristics,
whose inheritance did appear to be of a blending nature, could be
explained as resulting from the action of several different
Mendelian hereditary factors.

In the late 19th century improved microscopes and new stains

from the chemical industry had improved visualization of cells
and their nuclei. The German anatomist Walther Flemming,
working at Kiel, first identified chromosomes, and described the
process of cell division, now called mitosis, in which the
chromosomes enter the nuclei of both daughter cells. The Belgian

Genes before 1944

cytologist Edouard van Beneden showed that there was a
characteristic chromosome number for each species and that this
number was found in the various different cell types of an
organism. He followed chromosomes of the nematode worm
Ascaris through cell division and showed that the number was
conserved, but that it halved during the formation of the
reproductive cells (divisions forming reproductive cells are now
called ‘meioses’; singular, ‘meiosis’).

B the beginning of the 20th century, after Mendel’s work had been
resurrected, Theodor Boveri in Germany and Walter Sutton in the
USA independently showed that chromosomes behaved just like
Mendel’s hereditary factors. From then on most scientists believed
that the chromosomes were the hereditary factors or at least
contained them. The hereditary factors themselves were named
‘Gene’ (in German this is plural, equivalent to ‘genes’ in English)
in 1909 by Wilhelm Johannsen, Professor of Plant Physiology at
the University of Copenhagen. The term was derived from the
Greek γεѵεα (= generation or race). This is an interesting linguistic
example of the entity being named after the process, as ‘genetic’
7
 had been in use since 1830, and ‘genetics’, as a noun, was
introduced by William Bateson in 1905.

The centre of activity then moved to the USA where Thomas Hunt
Morgan, at Columbia University, effectively established modern
genetics through his studies of the fruit fly Drosophila. Drosophila
is a small insect with a short generation time. Large numbers can
be kept in small tubes, many crosses can be carried out in a
reasonable time scale, and insects have a lot of complex anatomy
on the surface that is easy to observe by simple visual examination.
Drosophila is therefore very well suited to genetic work in the
laboratory. Despite much effort, no treatments were found in the
early work that induced new mutations, but a number of
spontaneous mutations were discovered and used for breeding
studies. Whereas Mendel’s factors had separated from one another
in terms of their presence in individuals of the next generation,
many of Morgan’s Drosophila mutations did not do so. The
reason is that Drosophila has only four chromosomes, so it is
Genes

quite likely that two genes will lie on the same chromosome and
thus be transmitted together to reproductive cells. However,
genes on the same chromosome do sometimes separate and
Morgan was able to work out that this occurred because the
two parental chromosomes always form pairs with each other in
the last cell division (meiosis) leading to the formation of
reproductive cells (sperm or eggs). During this phase of pairing
the chromosomes can break and rejoin such that segments are
exchanged (see Figure 7 below). So a gene that was on one
parental chromosome can join one that was on the other, and be
transmitted together into the reproductive cells. Importantly
the probability of separation of two genes depends on the
distance between them and the frequency of segregation of
genes on the same chromosome became the basis for genetic
mapping. Using these techniques Morgan’s group created
accurate maps of the Drosophila chromosomes showing
the positions of many genes for which visible mutations
were available.
8
One of Morgan’s students, Hermann Muller, by then at the
University of Texas, finally discovered a treatment for Drosophila
that would generate new mutations. This was X-­irradiation. After
irradiation the chromosomes often showed visible changes,
strengthening the link between chromosomes and genes. In the
early 1930s it was found that the cells of the salivary glands of the
Drosophila larva contained giant chromosomes, much larger than
those found in most other cell types. These showed vastly more
detail down the microscope than normal-­size chromosomes and
when suitably stained they were seen to contain thousands of
bands, which looked like they might be single genes or small
groups of genes. As a result of this programme of work on
Drosophila, Morgan’s school had definitively proved that genes lay
on chromosomes, that they were arranged in a linear manner, and
that each cell contained a chromosome set from each parent such
that there were two copies of each in cells of the body and one

Genes before 1944

copy of each in reproductive cells.

In the late 19th century another scientific tradition had grown up

which we should now call statistics. In relation to evolution, a
school surrounding Karl Pearson (the ‘biometricians’) held to the
Darwinian orthodoxy and considered that inheritance was
inherently quantitative and continuous and that evolution
proceeded in small steps. Although apparently at variance with
Mendelism, the two traditions were eventually reconciled by the
work of R. A. Fisher. Fisher was for many years director of the
agricultural experimental station at Rothamsted, England, where
he devised most of the statistical methods now used for designing
and analysing quantitative experiments. He is also the father of
modern quantitative genetics. Fisher showed that quantitative
characters, such as height and weight, could be accounted for by
the action of several Mendelian genes, each with variants of
different effect, and that the blending behaviour seen in such cases
was completely consistent with Mendelian genetics. With Sewall
Wright, of the University of Chicago, and J. B. S. Haldane, then at
Cambridge University, Fisher created what has since been known
9
 as ‘the modern synthesis’, and also as Neo-­Darwinism, which is
essentially the theory of natural selection presented in terms of
Mendelian genetics.

During the early 20th century there was great parallel increase of
understanding about how living organisms transformed foodstuffs
into energy and into the materials for growth: what we now call
metabolism. It had been shown that each step in a metabolic
pathway was one chemical transformation catalysed by an enzyme
and that enzymes themselves were proteins. George Beadle and
Edward Tatum, working at Stanford University, chose the bread
mould Neurospora crassa as more amenable than Drosophila for
biochemical studies. Neurospora can be grown on simple agar
plates, in which the agar both acts as a surface for growth and also
contains the nutrient medium. If a mutation causes the inability
to synthesize a particular substance, the mould can still grow so
long as this substance is added to the medium. Beadle and Tatum
created many mutant strains by X-­irradiation and showed that
Genes

each mutation led to the lack of a specific enzyme. They concluded

that genes either were enzymes, or somehow controlled their
formation. This was later encapsulated in the slogan ‘one gene,
one enzyme’.

By the beginning of the Second World War a lot was known about
genes. It was understood that they lay on chromosomes in the cell
nucleus, that each chromosome came as a pair, with one member
from each parent, and that each gene somehow was responsible
for the production or activity of a specific enzyme. It was known
that changes in genes, mutations, could be produced by X-­rays,
and also by certain chemical substances which were therefore
called ‘mutagens’. What was not known was the chemical nature
of the gene. What were genes actually made of? When Addison
Gulick wrote his reviews this was a major issue. Like most others
at the time he presumed that they were large molecules, probably
consisting of proteins or protein complexes. But the identity of the

10
molecules, how they could be replicated, and how they could
control the formation of enzymes, remained a complete mystery.

DNA
The second line of work leading to understanding of the gene was
work on deoxyribonucleic acid, DNA. Contrary to general belief,
DNA has been known to biochemistry for a long time. It was
discovered in 1869 by Friedrich Miescher, a Swiss physiological
chemist working at the University of Tübingen in Germany.
Miescher studied white blood cells which he obtained in large
quantities by squeezing the pus out of the bandages of surgical
patients. He isolated nuclei from the cells using the enzyme pepsin,
then extracted them with weak alkali and precipitated with acid.
The resultant substance, called nuclein, was found to contain
phosphorus, but was quite unlike the other biological substances

Genes before 1944

known at the time to contain phosphorus. At the time, Miescher
considered the function of nuclein to be simply a store for
phosphorus within the body. Subsequent progress was quite slow by
modern standards. The reason for this is that, in contrast to biology,
where a breeding experiment could be conducted almost as easily in
1880 as 1980, in chemistry the techniques of the late 19th century
were very primitive. There were few methods for separating
substances, differential precipitation with salts or solvents being the
major technique. There were also few methods for analysing even
simple molecular structures, the mainstay being the purification of
the substance followed by a determination of the proportions by
weight of each chemical element that it contained. The difficulties
were particularly severe in biological chemistry, which deals with
large molecules that are easily damaged by harsh conditions.

In the period 1880–1900, Albrecht Kossel working at the

University of Berlin conducted extensive studies on nuclein.
During this time the substance became known as ‘nucleic acid’
because of the acidity arising from its high content of phosphate

11
 groups. Kossel found that major constituents of nucleic acids were
the five bases: adenine, thymine, cytosine, guanosine, and uracil.
In chemistry a ‘base’ is an alkaline substance, and all of these
substances are weakly alkaline because of the amino (NH2) groups
that they contain. The nucleic acid bases are usually represented
by the single letter abbreviations: A, T, C, G, U. In the early 20th
century extensive further chemical studies were made by Phoebus
Levene at the Rockefeller Institute for Medical Research in New
York. He identified the sugar ribose, and later its close chemical
relative deoxyribose, as components of nucleic acids. Most
importantly, he worked out that nucleic acids were long molecules
in which the bases, sugars, and phosphates were joined together
in a chain. By 1935 the actual chemical structure of nucleic acids
was essentially finalized with the identification of the positions of
the base, and the attachments of the two phosphates on each
sugar molecule (Figure 3). Without going into details, it will be
appreciated that the potential number of ways of joining bases,
sugars, and phosphates together is huge, and the discovery of the
Genes

specific linkages used was a major advance in understanding the

structure.

In terms of chemical nomenclature, a nucleoside consists of a base

joined to a sugar, a nucleotide consists of a nucleoside joined to a
phosphate group, and a nucleic acid consists of several nucleotides
joined together, as shown in Figure 3. During the 1920s it had
become appreciated that there were two major classes of nucleic
acid. ‘Animal nucleic acid’ (now called deoxyribonucleic acid or
DNA) contained deoxyribose as the sugar and A, C, G, and T as
bases. ‘Plant nucleic acid’ (now called ribonucleic acid or RNA)
contained ribose as the sugar and A, C, G, and U as the bases. In
reality animals and plants both contain DNA and RNA, but DNA
is found mostly in cell nuclei and RNA in the surrounding
cytoplasm. The names arose because DNA had been generally
extracted from calf thymus, which contains a lot of cells with large
nuclei and little cytoplasm, while RNA was usually extracted from
yeast, which has a small nucleus and a lot of cytoplasm. By the end
12
 A
–
O
– 5'
O P O C
O
O

G
3'
O
– 5'
O P O C
O
O

T
3'
O
– 5'
O P O C
O
O

C
3'
O

Genes before 1944

– 5'
O P O C
O
O

G
3'
O
– 5'
O P O C O
O

3'
OH

3. DNA two-­dimensional structure. This shows a chemical structure

of five nucleotides only. Each nucleotide consists of the sugar which
contains a ring of four carbon and one oxygen atoms, attached to a
base, depicted here just as A, C, G, T, and to the adjacent sugar by the
fifth carbon atom and a phosphate group. 5ʹ and 3ʹ are numbers given
to the different carbon atoms of the sugar.

13
 of the 1930s it was understood that the ‘animal’ and ‘plant’
nomenclature was not appropriate.

So, by the time of Addison Gulick’s reviews, a very accurate

picture of the gene had been built up from biological studies, and
the chemistry of DNA and RNA had been largely worked out. So
why did nobody realize that genes were made of DNA? There were
two very good reasons. First, the most popular view about the
structure of DNA was that it was a tetranucleotide; in other words
each molecule consisted of one of each of the nucleotides
containing A, C, G, and T. This was believed because when DNA
was degraded, approximately equal amounts of each of the four
bases were recovered. Also, the early measurements of the
molecular weight of DNA indicated a value of about 1,500,
equivalent in size to four nucleotides joined together.
A tetranucleotide would be a rather simple structure compared to
the proteins, which were known to be much larger and more
complex molecules, and it seemed impossible for genetic
Genes

information to be stored in something as simple as a

tetranucleotide. On the biological side, chromosomes were known
to contain genes and they also contained DNA. However, when
cells in the course of division were viewed down the microscope
using stains that reacted specifically with DNA, the colour was lost
during the cell division process. This indicated—­incorrectly—­that
the DNA became degraded and resynthesized in every cell cycle,
hardly behaviour compatible with the known long-­term stability
of the genes.

The chemical nature of the gene

Some of the most exciting discoveries in the life sciences have
been of the type where a particular substance is identified as being
responsible for a biological phenomenon. In this category belong
hormones, vitamins, and enzymes. But perhaps none was quite so
dramatic as the discovery of the chemical nature of genes. Genes
are, after all, at the absolute centre of biology. Viewed with
14
hindsight, discovery may seem painfully slow, but the discovery of
anything is remarkably difficult at the time. After all, there was no
guarantee that any one class of substance did make up the genes.
Perhaps genes were made of many complex substances and it was
the organization of huge molecular complexes that was all
important. Perhaps, even if pure genes could be analysed in terms
of their constituent molecules there would be no clue as to how
they actually worked.

In any case, you cannot identify a substance without a means of

measuring its activity. In biochemistry this means you need a
suitable bioassay, a procedure in which biological activity can be
demonstrated and measured. In the 1930s there seemed no
obvious way in which a gene might be put into a Drosophila or a
Neurospora in order to change the properties of the organism. The
bioassay for genes would need to involve something much simpler.

Genes before 1944

A method for putting genes into organisms had actually already
been invented by Frederick Griffith, working at the British
Ministry of Health Pathological Laboratory. Griffith was studying
the bacterium Streptococcus pneumoniae, a major cause of human
pneumonia as well as many other infections. These bacteria come
in various subtypes. The ‘smooth’ forms have slimy capsules
around the cells that enable them to evade the immune system of
the host, while the ‘rough’ forms do not. Accordingly the smooth
forms are much more pathogenic. The names ‘smooth’ and ‘rough’
actually relate to the appearance of colonies of the bacteria when
grown on agar plates. The key experiment of Griffith, published in
1928, was to show that when mice were infected with rough
bacteria plus heat-­killed smooth ones, they would develop the
infection and die. Following this, it was possible to recover live,
true-­breeding, smooth bacteria from the carcass. This did not
occur with rough bacteria alone, or with heat-­killed smooth
bacteria alone. The implication was that genes responsible for the
formation of the smooth capsules had passed from the dead
bacteria to the live ones.
15
 The work was taken further by the lab of Oswald Avery and
various co-­workers at the Rockefeller Institute for Medical
Research, New York. An important step forward was to show that
the effect would work in vitro. Rough bacteria treated with
chemical extracts of the smooth type would indeed yield smooth
colonies on agar plates, so the phenomenon did not require intact
mice with all the complexities of infection and immunity that are
going on in a whole animal. Eventually the extracts from the
smooth bacteria were purified to a high degree and were shown to
conform to the chemical composition of DNA. Moreover, the
activity could be destroyed by specific DNA-­degrading enzymes,
but not by other treatments. The seminal paper, published in
1944, is very cautious in its interpretation, and it only speculates
about the identity of the ‘transforming principle’ with a gene.
It did not create any great sensation at the time and was only
moderately cited in other works. However, its conclusions had
been noted by two individuals who were to play a big role in
events to come. They were Francis Crick and James Watson, and
Genes

Avery’s paper convinced them that DNA was really, really

important stuff.

16
Chapter 2
Genes as DNA

The double helix

After 1944, and as a result of Avery’s experiments, it was known to
a small group of cognoscenti that at least one type of gene, found
in Streptococcus pneumoniae, consisted of deoxyribonucleic acid
(DNA). Over the next 20 years, a remarkable set of discoveries
established the overall shape of modern molecular biology and
this story has been core textbook material ever since. Best known
among these events was the discovery of the three-­dimensional
structure of DNA: the famous double helix.

The chemical studies of the previous hundred years had shown

that the structure of the DNA molecule consisted of a long chain
of the four nucleotides, A, T, C, and G, joined by phosphate
groups, as shown in Figure 3. But there was an important
aspect of the chemical structure that was still unknown: the
three-­dimensional arrangement of these components in space.
This was deduced by a combination of X-­ray diffraction studies
carried out by Maurice Wilkins and Rosalind Franklin of King’s
College London, and molecular model building by James Watson
and Francis Crick at the University of Cambridge. Their work was
recognized by the award of the Nobel Prize for Physiology/
Medicine in 1962 (Rosalind Franklin was not a prize winner due

17
 to her early death in 1958, but her achievement, although
undervalued at the time, is now considered seminal).

The model building work was greatly helped by a critical piece of

chemical information that had recently been discovered by Erwin
Chargaff at Columbia University. This was the fact that DNA
contained equal numbers of molecules of A and T, and equal
numbers of C and G. On the other hand, the ratio of A+T/C+G
varied with the organism from which the DNA was isolated.
Previously it had been thought that all four nucleotides occurred
in equal proportions, hence the tetranucleotide hypothesis, but
Chargaff ’s measurements were more accurate than those of the
1920s and 1930s and the ratios suggested that A molecules must
be in some way associated with Ts, and Cs associated with Gs.

Although molecular structure does not always explain function, it

turned out that the three-­dimensional structure of DNA did, in
fact, explain how the substance could act as the genetic material.
Genes

One molecule of DNA consists of two strands, lying in opposite

senses in a helical configuration. They are held together by
attractive forces (hydrogen bonds) between the bases, such that
A pairs with T and C pairs with G (Figure 4). This base pairing
was consistent with ‘Chargaff ’s rule’: A=T and C=G. Because
the structure consisted of two complementary strands this
immediately suggested that DNA could be replicated by
separating the two strands and assembling a new complementary
strand on each, such that a new A is inserted opposite every T and
a new C opposite every G. The drama of the events leading up to
the discovery of the structure of DNA is nicely recounted in James
Watson’s book The Double Helix (1968).

Evidence that DNA actually did replicate in the way proposed was
soon obtained. Prior to each cell division, the entire DNA content
of each chromosome becomes copied into two double helical
molecules, each containing one of the original strands and one
newly synthesized strand. When cells divide, one of the two
18
 (a)
C G

T A (b)
H
A T H H
H O H N N
C C
T H
C C C C N
G C H C N H N C
sugar
C G N C C N
A T
sugar O H
T A
G Thymine Adenine

A T
H
T A H
N O N
H C
G C
HC C C C
G C N
H C N H N C sugar

Genes as DNA
N C C N
A T
sugar O H N
T A
H
G C Cytosine Guanine
C G
A

AT

4. DNA three-­dimensional structure. (a) DNA is a double helix

consisting of two strands running in opposite directions. The strands
are held together by hydrogen bonds between the bases. (b) Base pairs
formed by A‒T and C‒G.

identical double stranded molecules goes to one of the daughter

cells and the other copy goes to the other daughter. Hence both
daughter cells have exactly the same genes as the original cell. The
replication mechanism of DNA explains the essential process
whereby genes can be transmitted to daughter cells on cell
division, and to subsequent generations by sexual reproduction.

19
 The DNA structure also explains gene function within the
organism in terms of protein synthesis. It was known that genes
were responsible for making enzymes and other proteins. Proteins
are large molecules consisting of chains of amino acids of which
there are 20 main types, most of which are likely to be present in
any given protein. How could a sequence of four types of unit—­A,
T, C, G—­control formation of a protein sequence composed of 20
types of unit? This became known as the problem of the genetic
code. DNA has four different kinds of base so there are 16 possible
permutations of two bases and 64 possible permutations of three
bases. From this calculation it was early reasoned that it would
need at least three bases to specify a particular one of the amino
acids. The actual chemical machinery for making protein in
accordance with the DNA sequence turned out to be very complex
and to involve several kinds of ribonucleic acid (RNA) (Figure 5).
First the gene, which is a sequence of nucleotides on one strand of
the DNA double helix, is copied into messenger RNA (mRNA) in
a process called transcription, which is carried out by a number of
Genes

enzymes and other proteins. During transcription the mRNA

strand is assembled by placing each of the four bases opposite its
complement in the DNA sequence. RNA is a single stranded
molecule and also has four bases except that uracil (U) is used in
place of the thymine (T) in DNA. During transcription U is
inserted opposite A in DNA, and the other pairs are as usual:
A goes opposite T, C opposite G, G opposite C. The mRNA then
moves from the cell nucleus, where the DNA is located, into the
cytoplasm, where it associates with ribosomes. These are complex
bodies each consisting of three or four molecules of ribosomal
RNA (rRNA) together with about 100 specific molecules of
protein. The rRNA of ribosomes does not have an informational
function like mRNA; rather it serves to hold the ribosome
together and to carry out some of its catalytic functions.

In order to assemble a protein, a number of amino acid molecules

need to be joined into a linear chain. First, individual amino acid
molecules in the cytoplasm become attached to transfer RNAs
20
 Nucleus

DNA

mRNA transcription

Transfer
Mature mRNA
RNA

Growing
protein chain
Transport to
cytoplasm

mRNA

Genes as DNA
Ribosome Cell membrane

5. Protein synthesis. First the sequence of the gene is copied into

mRNA. This moves to the cell cytoplasm and a protein chain is
assembled according to the sequence with one amino acid represented
by each of the triplets of three nucleotides. Each amino acid
(represented by a black dot) is brought in by a tRNA molecule that
bears an exposed group of three nucleotides complementary to the
triplet in the mRNA.

(tRNAs). The tRNAs each contain a three base sequence which

recognizes three bases of the mRNA by the usual complementarity
rules (A binds U; C binds G). There is one type of tRNA for each
triplet. Each tRNA molecule brings in one amino acid molecule
which becomes joined to the growing protein chain. Of the 64
permutations of A, C, U, and G, 61 are used to specify amino acids,
and are recognized by specific transfer RNAs. The other three are
used as terminators, to specify the end of the protein chain. Once
21
 this is reached, the protein is complete and detaches from the
ribosome. The relationship between the 64 base triplets in
messenger RNA and the 20 amino acids is known as the genetic
code. The code is ‘degenerate’ in that several triplets can specify
the same amino acid. Some amino acids have as many as six
triplets, others have four, two, or just one. The process of
assembling the amino acid sequence of the protein in accordance
with the nucleotide sequence of the mRNA is known as
translation.

The package consisting of the double helical structure of DNA, its

mode of replication, messenger RNA, and the genetic code,
comprises the intellectual foundation of modern molecular
biology. These facts were discovered by a relatively small group of
people in the 10 years following the discovery of the double helix.
Since the 1960s the structure of genes and the mechanisms of
gene activity and protein synthesis have been studied in enormous
detail by hundreds of thousands of people. The current textbook
Genes

picture is very complex and involves thousands of individual

molecular components. However the essence of the process
remains as described here.

The second generation of molecular biology involved its

conversion into a practical toolkit for analysing and modifying
living organisms. This occurred later, in the 1970s and 1980s, and
involved the introduction of a number of techniques that have
now become extremely powerful. They include molecular cloning,
whereby single genes can be isolated and expanded to a chemical
quantity suitable for practical applications. Genes can now be
introduced into organisms by a whole range of methods, often
based on the use of viruses that normally introduce genes into
bacteria or animal cells. Also in the molecular biology toolkit is
DNA sequencing, which is a suite of chemical techniques for
determining the order, or ‘sequence’, of A, T, C, and G in the
molecules of a DNA sample. DNA sequencing techniques, first
invented in 1977, have progressed dramatically. The machines
22
Another random document with
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 Ritarit vaikenivat.

Ruhtinas Michal oli yleisesti suosittu miehuutensa,

anteliaisuutensa ja isänmaallisuutensa vuoksi. Kun koko Liettua
joutui venäläisten valtaan, niin hän yksin piti puoliaan Nieswieczissä,
ruotsalaisten alkaessa sotansa hän ei suostunut Janusz-ruhtinaan
houkutuksiinkaan oli ensimmäisenä yhtymässä liittoutuneihin.
Senvuoksi häntä nyt kuunneltiin. Muuten kenties kukaan myöskään
ei halunnut olla huonoissa väleissä niin mahtavan herran kanssa. Oli
miten oli, mutta sapelit pistettiin taas tuppeen, vieläpä muutamat
upseerit, jotka olivat Radziwillin klienttejä, huusivat:

— Ottakaa hänet pois tataarilaisilta! Tuomitkoon valtakunta hänet!

Pakanat eivät saa häväistä jaloa verta!

— Niin, ottakaamme hänet pois tataarilaisilta! — toisti ruhtinas.

— Löydämme aina panttivankeja, ja lunnaat hän maksaa itse. Herra
Woynillowicz, komentakaa miehenne liikkeelle, ja ottakoot he hänet
väkisin, jos eivät muuten saa!

— Minä tarjoudun panttivangiksi tataarilaisille! — huudahti herra

Gnoinski.

Wolodyjowski oli tällä välin mennyt Kmicicin luo ja sanoi:

— Mitä olettekaan tehnyt! Hän pääsee nyt ehein nahoin tästä

jutusta!

Kmicic hypähti kuin haavoitettu tiikeri.

— Herra ruhtinas! — huusi hän. — Hän on minun vankini! Minä

säästin hänen henkensä, mutta ehdoilla, jotka hän kerettiläisen
evankeliuminsa kautta vannoi täyttävänsä, ja mieluummin kuolen
kuin sallin hänen päästä niiden käsistä, joiden huostaan olen hänet
uskonut, ennenkuin hän on kaikki täyttänyt!

Näin sanoen hän kannusti hevostaan, ja synnynnäinen kiivaus

näytti jo saavan hänessä vallan, sillä hänen kasvonsa vääristyivät,
sieraimet laajenivat ja silmät alkoivat salamoida.

Samassa Woynillowicz ratsasti hänen eteensä.

— Pois tieltä, herra Babinicz! — huudahti hän.

— Pois tieltä, herra Woynillowicz! — karjaisi Kmicic ja iski

sapelinsa lappeella tämän hevosta niin voimakkaasti, että ratsu
horjahti kuin luoti olisi siihen sattunut ja lysähti etujaloilleen.

Syntyi suuri melu upseerien keskuudessa, mutta Gosiewski

tunkeutui esille ja sanoi:

— Hiljaa, hyvät herrat! Teidän ruhtinaallinen korkeutenne!

Hetmanivaltani nojalla julistan, että herra Babiniczilla on oikeus
tähän vankiin, ja sen, joka tahtoo saada hänet pois tataarilaisten
käsistä, pitää antaa hänen voittajalleen takuu.

Ruhtinas Michal hillitsi kiihtymyksensä ja sanoi Kmicicille:

— Sanokaa, mitä tahdotte!

— Että hän täyttää sovitut ehdot, ennenkuin pääsee vapaaksi!

— Hän täyttää ne, kun pääsee vapaaksi.

— En usko!
 — Siinä tapauksessa vannon hänen puolestaan Pyhän Neitsyen
nimessä, jota tunnustan, ja ritarikunniani kautta, että kaikki ehdot
täytetään. Muussa tapauksessa voitte vaatia minulta hyvitystä.

— Se riittää minulle! — sanoi Kmicic. — Mutta herra Gnoinskin on

ruvettava panttivangiksi, sillä muuten tataarilaiset panevat vastaan.
Minä luotan kunniasanaan.

— Kiitän teitä, herra ritari! — vastasi ruhtinas. — Älkää myöskään

pelätkö, että hän heti pääsee vapaaksi! Annan hänet herra hetmanin
huostaan siihen asti kunnes kuninkaan tuomio lankeaa.

— Olkoon niin! — sanoi hetmani. Hän käski Woynillowiczia

nousemaan toisen hevosen selkään, koska entinen oli vielä iskusta
pökerryksissä. Sitten hän lähetti hänet ja herra Gnoiskin hakemaan
ruhtinasta.

Mutta asia ei ollut vieläkään helposti suoritettavissa. Vanki täytyi

ottaa väkisin, sillä Hassun-bey pani ankarasti vastaa, ja vasta kun
hän näki herra Gnoiskin ja hänelle oli luvattu lunnaiksi satatuhatta
taaleria, hän rauhoittui.

Illalla ruhtinas Boguslaw jo oli Gosiewskin teltassa. Häntä

hoidettiin huolellisesti. Kaksi lääkäriä oli yhtä mittaa hänen luonaan.
Molemmat vakuuttivat hänen paranevan, sillä häntä oli haavoitettu
vain sapelin kärjellä, ja haava oli vaaraton.

Wolodyjowski ei voinut antaa Kmicicille anteeksi sitä, että tämä oli

säästänyt ruhtinaan hengen. Vihoissaan hän vältti ystäväänsä koko
päivän. Illalla tuli Kmicic itse hänen telttaansa.
 ‒ Jumalan tähden! ‒ huudahti hänet nähtyään pieni ritari. ‒
Kaikkea muuta olisin pikemmin odottanut kuin sitä, että päästätte
hengissä tuon petturin!

‒ Kuulkaa minua, herra Michal, ennen kuin tuomitsette! ‒ vastasi

Kmicic synkästi. ‒ Minun jalkani oli hänen rintansa päällä ja sapelini
hänen kaulallaan, mutta tiedättekö mitä tuo petturi minulle sanoi?…
Että Oleńka on käsketty surmata Taurogissa, jos ruhtinas tapetaan.
Mitä saatoin minä onneton tehdä? Ostin Oleńkan hengen hänen
hengellään… Mitä saatoin tehdä… Kristuksen nimessä… Mitä
saatoin tehdä…

Herra Andrzej alkoi repiä tukkaansa ja polkea kiivaasti jalkaa,

muta
Wolodyjowski mietti vähän aikaa ja sanoi:

‒ Minä ymmärrän teidän tuskanne… mutta sentään… te, nähkääs,

päästitte isänmaan petturin, joka vastaisuudessa voi tuottaa
isänmaalle raskaita onnettomuuksia… Olette tänään tehnyt
isänmaalle suuria palveluksia, mutta lopussa te uhrasitte yhteisen
edun omien yksityisten etujenne takia.

— Entä mitä te itse olisitte tehnyt, jos teille olisi sanottu, että veitsi
on pantu neiti Anna Borzobohatan kaulalle?

Wolodyjowski alkoi kiivaasti kierrellä viiksiään.

— Minä en väitä olevani mikään esimerkki. Hm! Mitäkö tekisin?…

Mutta Skrzetuski, jolla on roomalaisen sielu, ei olisi säästänyt
ruhtinaan henkeä, ja olen varma siitä, että Jumala ei olisi sallinut sen
vuoksi viattoman veren joutua vuotamaan.
 — Älä rankaise minua, Jumala, raskaitten rikosteni, vaan Sinun
laupeutesi mukaan!… Mutta että minä olisin allekirjoittanut oman
armaani kuolemantuomion…

Kmicic peitti kasvonsa käsiinsä.

— Auttakaa minua, taivaan enkelit! En koskaan! En koskaan!

— Tehty mikä tehty! — sanoi Wolodyjowski. Herra Andrzej veti

poveltaan esille papereita.

— Katsokaa, herra Michal, mitä olen saanut hänet antamaan!

Tässä on määräys Sakowiczille, tässä kaikille Radziwillin upseereille
ja ruotsalaisille komentajille… Hänen oli pakko allekirjoittaa, vaikka
tuskin jaksoi liikuttaa kättään… Ruhtinas Michal itse pakotti hänet
siihen… Tässä on Oleńkalle vapaus ja turvallisuus! Kautta Jumalan!
Minä ruoskin joka päivä itseäni, minä rakennutan uuden kirkon,
mutta hänen henkeään minä en uhraa! Minulla ei ole roomalaisen
sielua… olkoon niin! Minä en ole mikään Cato, kuten Skrzetuski…
myönnän sen! Mutta Oleńkaa en uhraa, en vaikka joutuisin alimpaan
helvettiin!…

Kmicic ei saanut lopetetuksi puhettaan, sillä Wolodyjowski riensi

hänen luokseen ja pannen kätensä hänen suulleen huusi
pelästyneenä:

— Älkää herjatko! Vedätte Jumalan rangaistuksen hänenkin

päälleen!
Lyökää rintaanne! Pian! Pian!

Kmicic alkoi lyödä rintaansa toistellen: "Mea culpa! Mea culpa!

Mea maxima culpa!" Viimein mies parka purskahti katkeraan itkuun,
sillä hän ei enää tietänyt mitä tekisi.

Wolodyjowski antoi hänen itkeä tarpeekseen. Viimein, kun Kmicic

oli rauhoittunut, hän kysyi häneltä:

— Mitä aiotte nyt tehdä?

— Lähden joukkoineni minne vain minut lähetetään, vaikkapa

Birzeen, kunhan vain miehet ja hevoset levähtävät. Matkan varrella
vuodatan Jumalan kunniaksi niin paljon kerettiläisten verta kuin
mahdollista.

— Se lasketaan teille ansioksi. Älkää masentuko! Jumala on

armollinen.

— Menen suorinta tietä! Koko Preussi on nyt avoinna, enintään

siellä täällä on jokin vähäinen varusväki.

Herra Michal huokasi.

— Mieleni tekee mukaanne aivan kuin paratiisiin. Mutta minun on

jäätävä hoitamaan tehtäviäni. Onnellinen olette te, jolla on
johdettavana vapaaehtoisia… Kuulkaahan, veliseni!… Kun tapaatte
ne molemmat, niin… pitäkäähän huolta siitä toisestakin, ettei hänelle
tapahtuisi mitään pahaa!… Jumala tietää, ehkäpä hänet on suotu
minulle…

Näin sanoen pieni ritari syleili Kmiciciä.

KAHDESTOISTA LUKU.

Oleńka ja Anusia poistuivat Braunin turvin Taurogista ja saapuivat

onnellisesti miekankantajan joukon luo, joka silloin oli Olszan luona
lähellä Taurogia.

Kun vanha aatelismies näki heidät molemmat hengissä ja

terveinä, hän ei ensin ollut uskoa silmiään, mutta alkoi sitten itkeä
ilosta ja tuli niin sotaiselle tuulelle, että ei enää pelännyt mitään
vaaroja. Tulkoonpa ei vain Boguslaw, vaan itse Ruotsin kuningaskin
koko sotavoimansa kera, niin miekankantaja kyllä oli valmis
suojelemaan tyttöjään!

— Ennen kaadun kuin hiuskarvakaan päästänne putoaa! — sanoi

hän. — En ole enää sama mies, jonka te tunsitte Taurogissa, ja
luulenpa ruotsalaisten kauan vielä muistavan tekojani. Totta on, että
petturi Sakowicz yllättäen hyökkäsi kimppuumme ja löi meidät, mutta
käytettävissäni on taas useita satoja sapeleita.

Miekankantaja oli todellakin suuresti muuttunut. Vaikea oli tuntea

häntä entiseksi masentuneeksi Taurogin vangiksi. Hän oli täynnä
tarmoa. Kedolla hevosen selässä hän oli omassa elementissään, ja,
hyvä sotilas kun oli, hän oli jo tuottanut ruotsalaisille paljon vahinkoa.
Koska hän oli hyvin arvossapidetty mies paikkakunnalla, kokoontui
hänen ympärilleen sekä aatelia että talonpoikia.

Miekankantajan joukkoon kuului kolmesataa talonpoikaa

jalkaväkeä ja noin viisisataa ratsumiestä. Vain harvoilla jalkamiehistä
oli tuliaseita. Useimmilla oli aseina viikatteet ja hangot. Aatelin
asestus oli parempaa, mutta hyvin kirjavaa sekin. Monella oli
humalaseipäät keihäitten asemesta. Toisilla oli isiltä perityt, upeat
aseet, jotka enimmäkseen olivat aivan vanhanaikaiset. Hevoset
olivat myös hyvin erilaisia.

Tämmöisellä joukolla saattoi miekankantaja sulkea tien

ruotsalaisilta patrulleilta, vieläpä tuhota suurempia ratsuväkiosastoja
sekä puhdistaa metsät ja kylät rosvoilevista joukoista, mutta hän ei
kyennyt käymään minkään kaupungin kimppuun.

Ruotsalaiset olivat viisastuneet vastoinkäymisistä. Heti kapinan

alettua oli kaikkialla Liettuassa ja Samogitiassa tuhottu pienemmät
joukko-osastot, jotka olivat hajaantuneina kyliin. Nyt jäljelläolevat
pysyttelivät linnoitetuissa kaupungeissa ja tekivät niistä retkiä vain
läheisyyteen. Tästä seurasi, että kedot, metsät, kylät ja
pikkukaupungit olivat puolalaisten käsissä, mutta kaikki suuremmat
kaupungit olivat ruotsalaisten hallussa, eikä heitä voitu niistä
mitenkään karkoittaa.

Miekankantajan "puolue" oli laatuaan parhaita. Muut saivat vielä

vähemmän toimeen. Liivinmaan rajalla olivat kapinoitsijat tosin
menneet uhkarohkeudessa niin pitkälle, että olivat kahdesti
piirittäneet Birzeä, jonka oli toisella kertaa täytynyt antautua, mutta
tämä tilapäinen menestys johtui siitä, että De la Gardie oli vienyt
kaikki joukkonsa puolustamaan Riikaa tsaarin hyökkäykseltä.
 Hänen harvinaisen loistavat voittonsa saattoivat kuitenkin
odottamaan, että sota siellä pian päättyy, ja sen jälkeen tulisivat
voittoisat ruotsalaiset sotajoukot taas Samogitiaan. Sillä aikaa
saattoivat kuitenkin lukuisat kapinoitsijain joukot olla turvallisina
metsissä varmoina siitä, että vihollinen ei tule heitä etsimään salojen
suojasta.

Näin ollen miekankantaja oli luopunut ajatuksesta etsiä turvaa

Bialowiezen korvesta, jonne oli pitkä matka ja matkan varrella monta
suurta kaupunkia, joissa oli vahva varusväki.

— Jumala on antanut meille kauniin syksyn, — puhui hän

tytöilleen, — ja siksi käy helpommin päinsä elää sub Jove. Annan
laittaa teille mukavan teltan ja hankin jonkun naisen teitä
palvelemaan, niin että voitte jäädä leiriin. Ei ole nykyaikoina
turvallisempaa olinpaikkaa kuin metsät. Minun Billewiczini on
porona, ja siellä kuljeksii sekä rosvoja että ruotsalaisia. Missäpä
teillä olisi turvallisempaa kuin minun luonani, jolla on käytettävänä
satoja sapeleita? Jos taas myöhemmin tulee sateita, niin löytyy teille
kyllä jokin maja korvessa.

Tämä ajatus miellytti suuresti neiti Borzobohataa, sillä joukossa oli

muutamia nuoria Billewiczejä, sieviä ritareita, ja sitäpaitsi huhuttiin
edelleen, että Babinicz oli tulossa sinne.

Anusia toivoi, että Babinicz saavuttuaan heti tekee lopun

ruotsalaisista, ja sitten… sitten käyköön niinkuin Jumala tahtoo.
Oleńkakin oli sitä mieltä, että turvallisinta oli leirissä, mutta hän tahtoi
olla mahdollisimman kaukana Taurogista, koska pelkäsi Sakowiczin
vainoa.
 — Menkäämme Wodoktyyn, — sanoi hän, — siellä olemme omien
ihmisten joukossa. Jos Wodokty on poltettu, on jäljellä Mitruny ja
kaikki ympäristön aateliskylät. On mahdotonta, että koko seutu olisi
autiona. Vaaran tullen suojelee Lauda meitä.

— Kaikki laudalaiset ovat lähteneet herra Wolodyjowskin mukaan,

— huomautti nuori Jur Billewicz.

— Vanhukset ja poikaset ovat jääneet jäljelle, ja sitäpaitsi naisetkin

siellä osaavat käyttää aseita. Myös salot ovat siellä suuremmat.
Domaszewiczit tai Gosiewiczit opastavat meidät Rogowskin
korpeen, josta meitä ei kukaan löydä.

— Ja minä saatettuani teidät turvaan käyn ruotsalaisten kimppuun

ja teen lopun niistä, jotka uskaltavat lähestyä salon reunaa, — sanoi
miekankantaja. — Tuo on oivallinen ajatus! Täällä meillä ei ole
mitään tekemistä, siellä voi saada aikaan paljon enemmän.

Kenpä tietää, eikö miekankantaja senvuoksi niin mieltynyt Oleńkan

ehdotukseen, että hänkin kaikessa hiljaisuudessa pelkäsi
Sakowiczia, joka nyt epätoivoissaan saattoi olla hirmuinen.

Neuvo oli muuten itsessäänkin hyvä ja miellytti senvuoksi kaikkia.

Vielä samana päivänä lähetti miekankantaja Jur Billewiczin johdolla
jalkaväen pyrkimään metsien läpi Krakinowiin. Itse hän lähti kaksi
päivää sen jälkeen ratsuväen kanssa liikkeelle otettuaan ennen sitä
selville, että Kiejdanyn ja Rosielnin välillä, mistä hänen oli kuljettava,
ei ollut ruotsalaista sotaväkeä.

He kulkivat hitaasti ja varovasti. Neidit ajoivat talonpoikaisrattailla

ja ratsastivat välillä miekankantajan heille hankkimilla hevosilla.
 Anusia, joka oli saanut Jurilta lahjaksi pienen sapelin, sitoi sen
vyölleen ja ratsasti kuin mikäkin ratsumestari joukon edessä. Hän
nautti retkestä ja auringonpaisteessa kimaltelevista sapeleista ja
öisistä leiritulista. Monet nuoret upseerit ja sotamiehet olivat
ihastuneet häneen, mutta hän heitti silmäyksiä joka puolelle, päästi
kulkiessa suortuvansa valloilleen ja sitten kolmasti päivässä palmikoi
ne taas käyttäen kuvastimenaan kirkasta lähdettä. Usein hän sanoi
haluavansa nähdä taistelun ja osoittaa rohkeuttaan, mutta se ei ollut
totta. Hän tyytyi vain valloittamaan kaikkien nuorten soturien
sydämet.

Myöskin Oleńka oli ikäänkuin uudelleen elpynyt paettuaan

Taurogista. Siellä häntä oli ahdistanut ainainen epävarmuus ja pelko,
mutta täällä metsän sydämessä hän tunsi olevansa turvassa. Raikas
ilma vahvisti häntä. Sotilaiden, aseiden ja leirielämän näkeminen
vaikuttivat kuin balsami hänen kiusattuun sieluunsa. Hänkin nautti
retkestä, eivätkä vaarat häntä ollenkaan peloittaneet, sillä hänen
suonissaan virtaili sotilasverta. Hän ei pyrkinyt kulkemaan joukon
etunenässä, minkä vuoksi hän myös vähemmän veti katseita
puoleensa. Mutta sen sijaan hän nautti yleistä kunnioitusta.

Kaikkien sotilaitten partaisille kasvoille levisi hymy, kun he näkivät

Anusian. Kun Oleńka lähestyi nuotioita, paljastuivat päät. Tämä
kunnioitus muuttui myöhemmin ihailuksi. Oli niitäkin, joiden nuoressa
rinnassa sydän sykki hänelle, mutta silmät eivät uskaltaneet katsoa
häneen yhtä rohkeasti kuin mustakiharaiseen ukrainattareen.

Kuljettiin läpi metsien ja viidakoiden ja lähetettiin usein edelle

tiedustelijoita, ja vihdoin myöhään seitsemännen päivän illalla
saavuttiin Lubicziin, joka oli Laudan rajalla ja ikäänkuin sen porttina.
Hevoset olivat sinä päivänä niin väsyneet, että Oleńkan esityksistä
huolimatta ei voitu jatkaa matkaa, vaan miekankantaja yöpyi sinne
joukkoineen. Hän itse neitien kanssa asettui taloon, sillä ilma oli
sumuinen ja kylmä. Omituisen sattuman kautta talo ei ollut joutunut
tulen uhriksi. Vihollinen oli säästänyt sen luultavasti ruhtinas Janusz
Radziwillin käskystä, koska se oli Kmicicin, ja vaikka ruhtinas
myöhemmin oli saanut tietää Kmicicin luopuneen hänestä, hän ei
ollut muistanut tai ennättänyt antaa uusia määräyksiä. Kapinalliset
luulivat sitä Billewiczien omaisuudeksi, rosvot taas eivät uskaltaneet
sitä ryöstää, koska se oli niin lähellä Laudaa. Kaikki oli siellä
senvuoksi ennallaan.

Katkeraa surua ja harmia tuntien Oleńka astui tämän katon alle.

Hän tunsi täällä joka nurkan, mutta joka paikkaan liittyi muisto
Kmicicistä. Oleńka kulki ruokasalin läpi, jossa oli Billewiczien
muotokuvia ja metsäeläinten kalloja. Luotien puhkomat kallot
riippuivat vielä nauloissaan, sapeliniskujen repimät muotokuvat
katselivat ankarina seiniltä aivan kuin olisivat tahtoneet sanoa:
"Katso, tyttö, katso, jälkeläisemme, kuinka hänen kätensä on
häväissyt jo kauan sitten hautaan vaipuneitten esi-isiesi kuvia!"

Oleńka tunsi, että tässä häväistyssä talossa hän ei saa

hetkeksikään unta. Kaikkialla pimeissä nurkissa hän oli näkevinään
Kmicicin hurjia tovereita. Miten nopeasti olikaan tuo mies, jota hän
niin oli rakastanut, lipunut mielivallasta lain rikkomiseen, siitä yhä
suurempiin rikoksiin, Radziwillin palvelukseen ja petokseen, jonka
huippuna oli käden kohottaminen kuningasta, koko valtakunnan isää
vastaan.

Yö kului, mutta uni ei ottanut tullakseen onnettoman Oleńkan

silmiin. Hänen sydämensä kaikkia haavoja alkoi kirvellä. Häpeä sai
uudelleen hänen poskensa hehkumaan, silmistä ei nyt vuotanut
kyyneliä, mutta sydämen täytti niin määrätön murhe, että se ei ollut
mahtua tuohon sydänparkaan.

Mitä hän suri? Sitä, että asiat voisivat olla toisin, jos tuo mies olisi
toisenlainen, jos hänellä kaikessa hillittömyydessään ja
hurjuudessaan edes olisi rehellinen sydän, jos hänen
rikollisuudellaan olisi edes jokin määrä, jos olisi jokin raja, jonka yli
hän ei voisi mennä. Hänelle voisi sentään antaa niin paljon
anteeksi!…

Anusia huomasi ystävättärensä kärsimykset ja aavisti niiden syyn,

sillä vanha miekankantaja oli jo aikaisemmin salaa kertonut hänelle
koko asian. Koska hänellä oli hyvä sydän, hän meni Oleńkan luo,
kietoi kätensä hänen kaulaansa ja sanoi:

— Oleńka! Sinun on katkeraa olla tässä talossa… Oleńka ei

alussa tahtonut mitään sanoa, alkoi vain väristä kuin haavanlehti,
mutta viimein hän purskahti hurjasti ja epätoivoisesti itkemään. Hän
tarttui suonenvedontapaisesti Anusian käteen, painoi vaaleat
kiharansa hänen olkaansa vastaan, ja nyyhkytykset puistelivat
hänen ruumistaan. Anusia sai odottaa kauan, ennenkuin kohtaus
meni ohitse. Viimein, kun Oleńka oli jonkin verran rauhoittunut,
Anusia kuiskasi:

— Oleńka, rukoilkaamme hänen puolestaan! Mutta Oleńka peitti

kasvot käsiinsä.

— En… voi!… — sai hän vaivoin sanotuksi. Ja vähän ajan

kuluttua hän heittäen kuumeisin liikkein taaksepäin hiuksensa, jotka
olivat valuneet hänen otsalleen, alkoi puhua tukahtuneella äänellä:
 — Katsohan… en voi!… Sinä olet onnellinen!… Sinun Babiniczizi
on jalo, mainehikas… Jumalan… ja isänmaan edessä… Sinä olet
onnellinen! Mutta minun ei ole lupa rukoillakaan… Täällä on
kaikkialla verta… raunioita! Jospa hän ei edes olisi pettänyt
isänmaata eikä olisi tarjoutunut myymään kuningasta!… Olin jo
antanut anteeksi kaikki… Kiejdanyssa… sillä ajattelin… sillä rakastin
häntä… koko sydämestäni… Mutta nyt en voi… oi laupias Jumala!
Minä en voi!… Soisin olevani kuollut… ja hänenkin kuolleen!

— Voimme rukoilla jokaisen sielun puolesta, sillä Jumala on

armahtavaisempi kuin ihmiset ja tuntee vaikuttimet, joita ihmiset
usein eivät tiedä, — sanoi Anusia.

Näin sanoen hän polvistui rukoilemaan, ja Oleńka heittäytyi

pitkäkseen lattialle ja oli siinä asennossa aamuun asti.

Seuraavana aamuna levisi ympäristöön tieto, että miekankantaja

Billewicz oli tullut Laudaan. Kaikki, jotka kynnelle kykenivät, tulivat
häntä tervehtimään. Läheisistä metsistä tuli tutisevia vanhuksia ja
naisia lapset sylissä. Kahteen vuoteen ei täällä kukaan ollut kylvänyt
eikä niittänyt. Kylät olivat suurimmaksi osaksi porona ja autioina.
Väestö asusti metsissä. Asekuntoiset miehet olivat lähteneet
Wolodyjowskin mukaan tahi liittyneet partiojoukkoihin. Vain poikaset
vartioivat omaisuuden jäännöksiä.

Miekankantajaa tervehdittiin pelastajana ja ilokyynelin, sillä näitten

yksinkertaisten ihmisten mielestä hänen ja neidin paluu
kotipaikoilleen oli merkkinä sodan ja onnettomuuksien loppumisesta.
Alettiin palata kyliin ja ajaa sinne metsästä puoleksi villiintynyttä
karjaa. Tosin ruotsalaisia oli lähellä Poniewiczin vallien suojassa,
mutta heistä ei nyt välitetty, kun miekankantajan joukko oli tullut, ja
sitä voitiin sitäpaitsi tarpeen vaatiessa vahvistaa lisämiehillä.
 Aikoipa herra Tomasz hyökätä Poniewiezin kimppuun
puhdistaakseen koko seudun vihollisista. Hän odotti vain, että vielä
enemmän miehiä kerääntyisi hänen lippujensa alle, ja varsinkin hän
koetti saada jalkaväelleen riittävästi ampuma-aseita, joita oli suuri
määrä piilossa metsissä. Niitä odotellessaan hän tarkasteli seutua
kulkien kylästä kylään.

Mutta näky, joka hänet kohtasi, oli surullinen. Kaikkialla oli palon ja
hävityksen jälkiä. Suuri osa väestöä oli surmattu ja miehiltä hakattu
kädet pois.

Näin kauheasti oli sota hävittänyt näitä seutuja, tämmöiset

seuraukset oli tuottanut ruhtinas Janusz Radziwillin petturuus.

Mutta ennenkuin miekankantaja oli lopettanut tarkastusmatkansa,

saapui uusia, samalla kertaa iloisia ja kamalia uutisia, jotka nopeasti
kulkivat suusta suuhun.

Jur Billewicz, joka pienen joukon kanssa oli ollut tiedusteluretkellä

ja saanut vangiksi muutamia ruotsalaisia, sai ensimmäisenä tiedon
Prostkin taistelusta. Senjälkeen tuli uusia sanomia, jotka olivat toinen
toistaan satumaisempia.

— Herra Gosiewski, — kerrottiin, — on voittanut kreivi Waldeckin,

Israelin ja ruhtinas Boguslawin. Vihollisarmeija on kokonaan tuhottu
ja sen johtajat vangittu. Koko Preussi on tulessa.

Muutamia viikkoja myöhemmin oli uusi peloittava nimi yhtä mittaa

kaikkien huulilla: Babinicz.

— Babiniczin ansiota on pääasiallisesti tuo voitto Prostkin luona,

— puhuttiin koko Samogitiassa. — Babinicz voitti ja otti kiinni omin
käsin ruhtinas Boguslawin.

Ja edelleen:

— Babinicz polttaa vaaliruhtinaan Preussia ja kulkee kuin surman

enkeli
Samogitiaa kohti hävittäen kaikki maan tasalle.

Ja lopuksi:

— Babinicz on polttanut poroksi Taurogin. Sakowicz on paennut

häntä ja piileksii metsissä.

Viimeksimainittu tapahtumapaikka oli niin lähellä, että varsin pian

huomattiin uutinen täydelleen paikkansa pitäväksi.

Koko ajan, kun näitä uutisia saapui, eli Anusia kuin kuumeessa,
nauroi ja itki vuorotellen, polki jalkaa, kun joku ei uskonut, ja toisteli
kaikille, halusivatpa nämä kuulla tai ei:

— Minä tunnen herra Babiniczin! Hän vei minut Zamośćiesta herra

Sapiehan luo. Hän on suurin soturi maailmassa. En luule edes herra
Czarnieckin vetävän vertoja hänelle, Sapiehan joukossa
palvellessaan hän retken alussa kukisti ruhtinas Boguslawin. Olen
varma siitä, että juuri hän voitti ruhtinaan myös Prostkin luona. Hän
voittaa vaikka kymmenen Sakowiczin kaltaista… Kuukaudessa hän
varmasti on karkoittanut ruotsalaiset koko Samogitiasta.

Hänen ennustuksensa alkoi nopeasti toteutua. Ei ollut enää

epäilystäkään, että kuuluisa soturi Babinicz oli tulossa Taurogista
pohjoista kohti.
 Koltyniassa hän voitti eversti Baldonin ja tuhosi perin pohjin hänen
joukkonsa. Wornin luona hän voitti Telsziin kulkevan ruotsalaisen
jalkaväen, ja Telszin luona hän taisteli suuren ja voittoisan taistelun
everstejä Normania ja Hudensköldiä vastaan, jossa Hudensköld
kaatui ja Norman miestensä jäännösten kera pakeni aina Samogitian
rajoille asti. Telszistä Babinicz kulki Kurszaniin ajaen edellään
pienempiä ruotsalaisjoukkoja, jotka pakenivat suurempien joukkojen
turviin.

Taurogista ja Polagista aina Birzeen ja Wilkomierziin lensi voittajan

maine. Kerrottiin julmuuksista, joita hän harjoitti ruotsalaisia kohtaan.
Sanottiin, että hänen sotajoukkonsa, johon alussa oli kuulunut vain
pieni tataarilaisrykmentti ja jonkin verran vapaaehtoisia, kasvoi päivä
päivältä, sillä ken vain voi ja kaikki partiojoukotkin liittyivät häneen, ja
hän taivutti ne kaikki rautaisen tahtonsa alle ja johti vihollista
vastaan.

Huomio oli niin kiintynyt hänen voittoihinsa, että tieto tappiosta,

jonka Stenbock oli tuottanut Gosiewskille, Filipowin luona, jäi
melkein huomaamatta. Babinicz oli lähempänä, ja hänen toimiaan
kaikki seurasivat.

Anusia rukoili joka päivä miekankantajaa yhtymään

mainehikkaaseen soturiin. Oleńka kannatti häntä. Niin tekivät myös
kaikki upseerit ja aateli, joita uteliaisuuskin vaivasi.

Mutta tuuma ei ollut helppo toteuttaa. Ensiksikin Babinicz oli aivan

toisella seudulla. Toiseksi hän usein katosi viikkomääriksi
tietymättömiin, ja sitten taas yht'äkkiä kuultiin jotakin hänen uudesta
voitostaan. Kolmanneksi olivat tiet täynnä häntä pakenevia
ruotsalaisia. Vihdoin oli myös Sakowicz lähtenyt liikkeelle melkoisen
suuren joukon kanssa ja hävitti kauheasti kaikki tieltään sekä koetti
kiduttamalla saada ihmisiltä tietää, missä Billewiczien vapaajoukko
oli.

Kaiken lisäksi miekankantaja vielä pelkäsi, että olo Laudan

seuduilla käy pian hänelle liian vaikeaksi.

Tietämättä oikein itsekään, mitä oli tehtävä, hän uskoi Jur

Billewiczille aikovansa vetäytyä länteen päin saloille. Jur kertoi
tämän heti Anusialle, ja tämä meni oikopäätä miekankantajan
puheille.

— Rakas setä, — sanoi hän hänelle (näin hän nimitti häntä aina,
kun tahtoi saada hänet johonkin suostumaan), — olen kuullut, että
aiomme paeta. Eikö ole häpeä noin kuuluisalle soturille lähteä
pakoon heti, kun vain kuulee vihollisen olevan tulossa?

— Teidän täytyy pistää joka paikkaan nenänne! sanoi

miekankantaja loukkaantuneena. — Tämä ei ole teidän asianne!

— Hyvä! Jos te pakenette, niin minä jään!

— Jottako Sakowicz voisi teidät vangita?

— Sakowicz ei vangitse minua, sillä Babinicz puolustaa minua.

— Eihän hän tiedä, missä te olette! Olenhan sanonut, että emme

voi päästä hänen luokseen.

— Mutta hän voi tulla meidän luoksemme. Olen hänen tuttunsa.

Jos vain voisin lähettää hänelle kirjeen, niin olen varma siitä, että
hän tulisi heti, kun on voittanut Sakowiczin. Hän piti minusta hiukan,
ja siksipä hän ei kieltäisi apuaan.
 — Ja kuka vie hänelle kirjeen?

— Sitä viemään voi lähettää jonkun talonpojan.

— Kannattaahan joka tapauksessa koettaa. Oleńkalla on terävä

järki, mutta onpa järkeä teilläkin. Vaikka meidän sillä välin pitäisikin
piillä ylivoimaista vihollista metsässä, niin on kuitenkin hyvä, että
Babinicz tulee näille seuduille, sillä siten pääsemme pikemmin
häneen yhtymään. Yrittäkää! Luotettavan sanansaattajan kyllä
löydämme.

Anusia ryhtyi iloissaan toimimaan semmoisella menestyksellä, että

hänellä jo samana päivänä oli kaksikin kirjeenviejää, Jur Billewicz ja
Braun. Molempien oli määrä saada mukaansa samanlainen kirje,
jotta joka tapauksessa Babinicz saisi sanan. Enemmän puuhaa oli
Anusialla itse kirjeen sepittämisessä. Viimein hän laati sen näin
kuuluvaksi: "Suuressa vaarassa ollen kirjoitan Teille, jos ehkä
muistatte minut (vaikka mitä syytä minulla olisi luulla, että muistatte),
pyytäen Teitä tulemaan apuun. Muistaen sen hyväntahtoisuuden,
jota osoititte minulle matkalla Zamośćiesta, rohkenen toivoa, että
ette minua vaarassa hylkää. Olen herra miekankantaja Billewiczin
joukon turvissa. Hän on antanut minulle suojaa, koska olen auttanut
hänen sukulaisensa neiti Billewiczin pakoon vankeudesta Taurogista.
Sekä häntä että meitä ahdistaa joka puolelta vihollinen, nimittäin
ruotsalaiset sekä muuan herra Sakowicz, jonka tunkeilevaisuutta
paetakseni minun oli pakko hakea turvaa sotilasleiristä. Tiedän, että
Te ette minusta pitänyt, mutta Jumala näkee, että en ole tehnyt teille
mitään pahaa ja että aina koko sydämestäni olen ollut teille
myötätuntoinen. Mutta pelastakaa kuitenkin orpo tyttöraukka
vihollisen käsistä! Jumala palkitsee sen teille satakertaisesti, ja minä
rukoilen aina Teidän puolestanne, jota nyt nimitän vain hyväksi
suojelijakseni, mutta sitten pelastajakseni aina kuolemaani asti."

Kun lähettiläät jo olivat lähdössä, alkoi Anusia ajatella niitä

vaaroja, joihin he saattoivat joutua, ja koetti estää heitä lähtemästä.
Kyynelsilmin hän pyysi miekankantajaa kieltämään heitä lähtemästä
matkaan, sillä kirjeet saattoi yhtä hyvin antaa talonpoikien vietäväksi,
joiden oli helpompi päästä perille.

Mutta Braun ja Jur Billewicz pysyivät niin itsepintaisesti

päätöksessään lähteä, että heihin ei vaikuttanut mikään. Kilvan he
kumpikin tahtoivat osoittaa alttiuttaan, eikä kumpikaan aavistanut,
mikä heitä odotti. Viikkoa myöhemmin Braun joutui Sakowiczin
vangiksi, ja tämä antoi nylkeä häneltä nahan. Jur parka ammuttiin
viikkoa myöhemmin, kun hän Poniewiezen luona koetti paeta erästä
ruotsalaista joukkuetta.

Molemmat kirjeet joutuivat vihollisen käsiin.