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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 David A. Gerber
Charles L. Cohen AMERICAN INTELLECTUAL
ACCOUNTING Christopher Nobes HISTORY
ADDICTION Keith Humphreys Jennifer Ratner-­Rosenhagen
ADOLESCENCE Peter K. Smith THE AMERICAN JUDICIAL SYSTEM
THEODOR W. ADORNO Charles L. Zelden
Andrew Bowie AMERICAN LEGAL HISTORY
ADVERTISING Winston Fletcher G. Edward White
AERIAL WARFARE Frank Ledwidge AMERICAN MILITARY HISTORY
AESTHETICS Bence Nanay Joseph T. Glatthaar
AFRICAN AMERICAN HISTORY AMERICAN NAVAL HISTORY
Jonathan Scott Holloway Craig L. Symonds
AFRICAN AMERICAN RELIGION AMERICAN POETRY David Caplan
Eddie S. Glaude Jr AMERICAN POLITICAL HISTORY
AFRICAN HISTORY John Parker and Donald Critchlow
Richard Rathbone AMERICAN POLITICAL PARTIES
AFRICAN POLITICS Ian Taylor AND ELECTIONS L. Sandy Maisel
AFRICAN RELIGIONS AMERICAN POLITICS
Jacob K. Olupona Richard M. Valelly
AGEING Nancy A. Pachana THE AMERICAN PRESIDENCY
AGNOSTICISM Robin Le Poidevin Charles O. Jones
AGRICULTURE Paul Brassley and THE AMERICAN REVOLUTION
Richard Soffe Robert J. Allison
ALEXANDER THE GREAT AMERICAN SLAVERY
Hugh Bowden Heather Andrea Williams
ALGEBRA Peter M. Higgins THE AMERICAN SOUTH
AMERICAN BUSINESS HISTORY Charles Reagan Wilson
Walter A. Friedman THE AMERICAN WEST
AMERICAN CULTURAL HISTORY Stephen Aron
Eric Avila AMERICAN WOMEN’S HISTORY
AMERICAN FOREIGN RELATIONS Susan Ware
Andrew Preston AMPHIBIANS T. S. Kemp
AMERICAN HISTORY Paul S. Boyer ANAESTHESIA Aidan O’Donnell
ANALYTIC PHILOSOPHY ASTROPHYSICS James Binney
Michael Beaney ATHEISM Julian Baggini
ANARCHISM Alex Prichard THE ATMOSPHERE Paul I. Palmer
ANCIENT ASSYRIA Karen Radner AUGUSTINE Henry Chadwick
ANCIENT EGYPT Ian Shaw JANE AUSTEN Tom Keymer
ANCIENT EGYPTIAN ART AND AUSTRALIA Kenneth Morgan
ARCHITECTURE Christina Riggs AUTISM Uta Frith
ANCIENT GREECE Paul Cartledge AUTOBIOGRAPHY Laura Marcus
ANCIENT GREEK AND ROMAN THE AVANT GARDE
SCIENCE Liba Taub David Cottington
THE ANCIENT NEAR EAST THE AZTECS Davíd Carrasco
Amanda H. Podany BABYLONIA Trevor Bryce
ANCIENT PHILOSOPHY Julia Annas BACTERIA Sebastian G. B. Amyes
ANCIENT WARFARE BANKING John Goddard and
Harry Sidebottom John O. S. Wilson
ANGELS David Albert Jones BARTHES Jonathan Culler
ANGLICANISM Mark Chapman THE BEATS David Sterritt
THE ANGLO-SAXON AGE John Blair BEAUTY Roger Scruton
ANIMAL BEHAVIOUR LUDWIG VAN BEETHOVEN
Tristram D. Wyatt Mark Evan Bonds
THE ANIMAL KINGDOM BEHAVIOURAL ECONOMICS
Peter Holland Michelle Baddeley
ANIMAL RIGHTS David DeGrazia BESTSELLERS John Sutherland
ANSELM Thomas Williams THE BIBLE John Riches
THE ANTARCTIC Klaus Dodds BIBLICAL ARCHAEOLOGY
ANTHROPOCENE Erle C. Ellis Eric H. Cline
ANTISEMITISM Steven Beller BIG DATA Dawn E. Holmes
ANXIETY Daniel Freeman and BIOCHEMISTRY Mark Lorch
Jason Freeman BIOGEOGRAPHY Mark V. Lomolino
THE APOCRYPHAL BIOGRAPHY Hermione Lee
GOSPELS Paul Foster BIOMETRICS Michael Fairhurst
APPLIED MATHEMATICS ELIZABETH BISHOP
Alain Goriely Jonathan F. S. Post
THOMAS AQUINAS Fergus Kerr BLACK HOLES Katherine Blundell
ARBITRATION Thomas Schultz and BLASPHEMY Yvonne Sherwood
Thomas Grant BLOOD Chris Cooper
ARCHAEOLOGY Paul Bahn THE BLUES Elijah Wald
ARCHITECTURE Andrew Ballantyne THE BODY Chris Shilling
THE ARCTIC Klaus Dodds and THE BOHEMIANS David Weir
Jamie Woodward NIELS BOHR J. L. Heilbron
HANNAH ARENDT Dana Villa THE BOOK OF COMMON PRAYER
ARISTOCRACY William Doyle Brian Cummings
ARISTOTLE Jonathan Barnes THE BOOK OF MORMON
ART HISTORY Dana Arnold Terryl Givens
ART THEORY Cynthia Freeland BORDERS Alexander C. Diener and
ARTIFICIAL INTELLIGENCE Joshua Hagen
Margaret A. Boden THE BRAIN Michael O'Shea
ASIAN AMERICAN HISTORY BRANDING Robert Jones
Madeline Y. Hsu THE BRICS Andrew F. Cooper
ASTROBIOLOGY David C. Catling BRITISH CINEMA Charles Barr
THE BRITISH CONSTITUTION COGNITIVE BEHAVIOURAL
Martin Loughlin THERAPY Freda McManus
THE BRITISH EMPIRE Ashley Jackson COGNITIVE NEUROSCIENCE
BRITISH POLITICS Tony Wright Richard Passingham
BUDDHA Michael Carrithers THE COLD WAR Robert J. McMahon
BUDDHISM Damien Keown COLONIAL AMERICA Alan Taylor
BUDDHIST ETHICS Damien Keown COLONIAL LATIN AMERICAN
BYZANTIUM Peter Sarris LITERATURE Rolena Adorno
CALVINISM Jon Balserak COMBINATORICS Robin Wilson
ALBERT CAMUS Oliver Gloag COMEDY Matthew Bevis
CANADA Donald Wright COMMUNISM Leslie Holmes
CANCER Nicholas James COMPARATIVE LITERATURE
CAPITALISM James Fulcher Ben Hutchinson
CATHOLICISM Gerald O’Collins COMPETITION AND ANTITRUST
CAUSATION Stephen Mumford and LAW Ariel Ezrachi
Rani Lill Anjum COMPLEXITY John H. Holland
THE CELL Terence Allen and THE COMPUTER Darrel Ince
Graham Cowling COMPUTER SCIENCE
THE CELTS Barry Cunliffe Subrata Dasgupta
CHAOS Leonard Smith CONCENTRATION CAMPS
GEOFFREY CHAUCER David Wallace Dan Stone
CHEMISTRY Peter Atkins CONDENSED MATTER PHYSICS
CHILD PSYCHOLOGY Usha Goswami Ross H. McKenzie
CHILDREN’S LITERATURE CONFUCIANISM Daniel K. Gardner
Kimberley Reynolds THE CONQUISTADORS
CHINESE LITERATURE Sabina Knight Matthew Restall and
CHOICE THEORY Michael Allingham Felipe Fernández-Armesto
CHRISTIAN ART Beth Williamson CONSCIENCE Paul Strohm
CHRISTIAN ETHICS D. Stephen Long CONSCIOUSNESS Susan Blackmore
CHRISTIANITY Linda Woodhead CONTEMPORARY ART
CIRCADIAN RHYTHMS Julian Stallabrass
Russell Foster and Leon Kreitzman CONTEMPORARY FICTION
CITIZENSHIP Richard Bellamy Robert Eaglestone
CITY PLANNING Carl Abbott CONTINENTAL PHILOSOPHY
CIVIL ENGINEERING Simon Critchley
David Muir Wood COPERNICUS Owen Gingerich
THE CIVIL RIGHTS MOVEMENT CORAL REEFS Charles Sheppard
Thomas C. Holt CORPORATE SOCIAL
CLASSICAL LITERATURE RESPONSIBILITY Jeremy Moon
William Allan CORRUPTION Leslie Holmes
CLASSICAL MYTHOLOGY COSMOLOGY Peter Coles
Helen Morales COUNTRY MUSIC Richard Carlin
CLASSICS Mary Beard and CREATIVITY Vlad Glăveanu
John Henderson CRIME FICTION Richard Bradford
CLAUSEWITZ Michael Howard CRIMINAL JUSTICE
CLIMATE Mark Maslin Julian V. Roberts
CLIMATE CHANGE Mark Maslin CRIMINOLOGY Tim Newburn
CLINICAL PSYCHOLOGY CRITICAL THEORY
Susan Llewelyn and Katie Stephen Eric Bronner
Aafjes-van Doorn THE CRUSADES Christopher Tyerman
CRYPTOGRAPHY Fred Piper and EMOTION Dylan Evans
Sean Murphy EMPIRE Stephen Howe
CRYSTALLOGRAPHY A. M. Glazer EMPLOYMENT LAW David Cabrelli
THE CULTURAL REVOLUTION ENERGY SYSTEMS Nick Jenkins
Richard Curt Kraus ENGELS Terrell Carver
DADA AND SURREALISM ENGINEERING David Blockley
David Hopkins THE ENGLISH LANGUAGE
DANTE Peter Hainsworth and Simon Horobin
David Robey ENGLISH LITERATURE Jonathan Bate
DARWIN Jonathan Howard THE ENLIGHTENMENT
THE DEAD SEA SCROLLS John Robertson
Timothy H. Lim ENTREPRENEURSHIP Paul Westhead
DECADENCE David Weir and Mike Wright
DECOLONIZATION Dane Kennedy ENVIRONMENTAL
DEMENTIA Kathleen Taylor ECONOMICS Stephen Smith
DEMOCRACY Naomi Zack ENVIRONMENTAL ETHICS
DEMOGRAPHY Sarah Harper Robin Attfield
DEPRESSION Jan Scott and ENVIRONMENTAL LAW
Mary Jane Tacchi Elizabeth Fisher
DERRIDA Simon Glendinning ENVIRONMENTAL POLITICS
DESCARTES Tom Sorell Andrew Dobson
DESERTS Nick Middleton ENZYMES Paul Engel
DESIGN John Heskett EPICUREANISM Catherine Wilson
DEVELOPMENT Ian Goldin EPIDEMIOLOGY Rodolfo Saracci
DEVELOPMENTAL BIOLOGY ETHICS Simon Blackburn
Lewis Wolpert ETHNOMUSICOLOGY Timothy Rice
THE DEVIL Darren Oldridge THE ETRUSCANS Christopher Smith
DIASPORA Kevin Kenny EUGENICS Philippa Levine
CHARLES DICKENS Jenny Hartley THE EUROPEAN UNION
DICTIONARIES Lynda Mugglestone Simon Usherwood and John Pinder
DINOSAURS David Norman EUROPEAN UNION LAW
DIPLOMATIC HISTORY Anthony Arnull
Joseph M. Siracusa EVANGELICALISM
DOCUMENTARY FILM John G. Stackhouse Jr.
Patricia Aufderheide EVIL Luke Russell
DREAMING J. Allan Hobson EVOLUTION Brian and
DRUGS Les Iversen Deborah Charlesworth
DRUIDS Barry Cunliffe EXISTENTIALISM Thomas Flynn
DYNASTY Jeroen Duindam EXPLORATION Stewart A. Weaver
DYSLEXIA Margaret J. Snowling EXTINCTION Paul B. Wignall
EARLY MUSIC Thomas Forrest Kelly THE EYE Michael Land
THE EARTH Martin Redfern FAIRY TALE Marina Warner
EARTH SYSTEM SCIENCE Tim Lenton FAMILY LAW Jonathan Herring
ECOLOGY Jaboury Ghazoul MICHAEL FARADAY
ECONOMICS Partha Dasgupta Frank A. J. L. James
EDUCATION Gary Thomas FASCISM Kevin Passmore
EGYPTIAN MYTH Geraldine Pinch FASHION Rebecca Arnold
EIGHTEENTH‑CENTURY BRITAIN FEDERALISM Mark J. Rozell and
Paul Langford Clyde Wilcox
THE ELEMENTS Philip Ball FEMINISM Margaret Walters
FILM Michael Wood GLOBAL ECONOMIC HISTORY
FILM MUSIC Kathryn Kalinak Robert C. Allen
FILM NOIR James Naremore GLOBAL ISLAM Nile Green
FIRE Andrew C. Scott GLOBALIZATION Manfred B. Steger
THE FIRST WORLD WAR GOD John Bowker
Michael Howard GÖDEL’S THEOREM A. W. Moore
FLUID MECHANICS Eric Lauga GOETHE Ritchie Robertson
FOLK MUSIC Mark Slobin THE GOTHIC Nick Groom
FOOD John Krebs GOVERNANCE Mark Bevir
FORENSIC PSYCHOLOGY GRAVITY Timothy Clifton
David Canter THE GREAT DEPRESSION AND
FORENSIC SCIENCE Jim Fraser THE NEW DEAL Eric Rauchway
FORESTS Jaboury Ghazoul HABEAS CORPUS Amanda L. Tyler
FOSSILS Keith Thomson HABERMAS James Gordon Finlayson
FOUCAULT Gary Gutting THE HABSBURG EMPIRE Martyn Rady
THE FOUNDING FATHERS HAPPINESS Daniel M. Haybron
R. B. Bernstein THE HARLEM RENAISSANCE
FRACTALS Kenneth Falconer Cheryl A. Wall
FREE SPEECH Nigel Warburton THE HEBREW BIBLE AS
FREE WILL Thomas Pink LITERATURE Tod Linafelt
FREEMASONRY Andreas Önnerfors HEGEL Peter Singer
FRENCH LITERATURE John D. Lyons HEIDEGGER Michael Inwood
FRENCH PHILOSOPHY THE HELLENISTIC AGE
Stephen Gaukroger and Knox Peden Peter Thonemann
THE FRENCH REVOLUTION HEREDITY John Waller
William Doyle HERMENEUTICS Jens Zimmermann
FREUD Anthony Storr HERODOTUS Jennifer T. Roberts
FUNDAMENTALISM Malise Ruthven HIEROGLYPHS Penelope Wilson
FUNGI Nicholas P. Money HINDUISM Kim Knott
THE FUTURE Jennifer M. Gidley HISTORY John H. Arnold
GALAXIES John Gribbin THE HISTORY OF ASTRONOMY
GALILEO Stillman Drake Michael Hoskin
GAME THEORY Ken Binmore THE HISTORY OF CHEMISTRY
GANDHI Bhikhu Parekh William H. Brock
GARDEN HISTORY Gordon Campbell THE HISTORY OF CHILDHOOD
GENES Jonathan Slack James Marten
GENIUS Andrew Robinson THE HISTORY OF CINEMA
GENOMICS John Archibald Geoffrey Nowell-Smith
GEOGRAPHY John Matthews and THE HISTORY OF COMPUTING
David Herbert Doron Swade
GEOLOGY Jan Zalasiewicz THE HISTORY OF EMOTIONS
GEOMETRY Maciej Dunajski Thomas Dixon
GEOPHYSICS William Lowrie THE HISTORY OF LIFE
GEOPOLITICS Klaus Dodds Michael Benton
GERMAN LITERATURE Nicholas Boyle THE HISTORY OF MATHEMATICS
GERMAN PHILOSOPHY Jacqueline Stedall
Andrew Bowie THE HISTORY OF MEDICINE
THE GHETTO Bryan Cheyette William Bynum
GLACIATION David J. A. Evans THE HISTORY OF PHYSICS
GLOBAL CATASTROPHES Bill McGuire J. L. Heilbron
THE HISTORY OF POLITICAL INSECTS Simon Leather
THOUGHT Richard Whatmore IRAN Ali M. Ansari
THE HISTORY OF TIME ISLAM Malise Ruthven
Leofranc Holford‑Strevens ISLAMIC HISTORY Adam Silverstein
HIV AND AIDS Alan Whiteside ISLAMIC LAW Mashood A. Baderin
HOBBES Richard Tuck ISOTOPES Rob Ellam
HOLLYWOOD Peter Decherney ITALIAN LITERATURE
THE HOLY ROMAN EMPIRE Peter Hainsworth and David Robey
Joachim Whaley HENRY JAMES Susan L. Mizruchi
HOME Michael Allen Fox JAPANESE LITERATURE Alan Tansman
HOMER Barbara Graziosi JESUS Richard Bauckham
HORMONES Martin Luck JEWISH HISTORY David N. Myers
HORROR Darryl Jones JEWISH LITERATURE Ilan Stavans
HUMAN ANATOMY Leslie Klenerman JOURNALISM Ian Hargreaves
HUMAN EVOLUTION Bernard Wood JAMES JOYCE Colin MacCabe
HUMAN PHYSIOLOGY JUDAISM Norman Solomon
Jamie A. Davies JUNG Anthony Stevens
HUMAN RESOURCE THE JURY Renée Lettow Lerner
MANAGEMENT Adrian Wilkinson KABBALAH Joseph Dan
HUMAN RIGHTS Andrew Clapham KAFKA Ritchie Robertson
HUMANISM Stephen Law KANT Roger Scruton
HUME James A. Harris KEYNES Robert Skidelsky
HUMOUR Noël Carroll KIERKEGAARD Patrick Gardiner
THE ICE AGE Jamie Woodward KNOWLEDGE Jennifer Nagel
IDENTITY Florian Coulmas THE KORAN Michael Cook
IDEOLOGY Michael Freeden KOREA Michael J. Seth
THE IMMUNE SYSTEM LAKES Warwick F. Vincent
Paul Klenerman LANDSCAPE ARCHITECTURE
INDIAN CINEMA Ian H. Thompson
Ashish Rajadhyaksha LANDSCAPES AND
INDIAN PHILOSOPHY Sue Hamilton GEOMORPHOLOGY
THE INDUSTRIAL REVOLUTION Andrew Goudie and Heather Viles
Robert C. Allen LANGUAGES Stephen R. Anderson
INFECTIOUS DISEASE Marta L. Wayne LATE ANTIQUITY Gillian Clark
and Benjamin M. Bolker LAW Raymond Wacks
INFINITY Ian Stewart THE LAWS OF THERMODYNAMICS
INFORMATION Luciano Floridi Peter Atkins
INNOVATION Mark Dodgson and LEADERSHIP Keith Grint
David Gann LEARNING Mark Haselgrove
INTELLECTUAL PROPERTY LEIBNIZ Maria Rosa Antognazza
Siva Vaidhyanathan C. S. LEWIS James Como
INTELLIGENCE Ian J. Deary LIBERALISM Michael Freeden
INTERNATIONAL LAW LIGHT Ian Walmsley
Vaughan Lowe LINCOLN Allen C. Guelzo
INTERNATIONAL MIGRATION LINGUISTICS Peter Matthews
Khalid Koser LITERARY THEORY Jonathan Culler
INTERNATIONAL RELATIONS LOCKE John Dunn
Christian Reus-Smit LOGIC Graham Priest
INTERNATIONAL SECURITY LOVE Ronald de Sousa
Christopher S. Browning MARTIN LUTHER Scott H. Hendrix
MACHIAVELLI Quentin Skinner JOHN STUART MILL Gregory Claeys
MADNESS Andrew Scull MINERALS David Vaughan
MAGIC Owen Davies MIRACLES Yujin Nagasawa
MAGNA CARTA Nicholas Vincent MODERN ARCHITECTURE
MAGNETISM Stephen Blundell Adam Sharr
MALTHUS Donald Winch MODERN ART David Cottington
MAMMALS T. S. Kemp MODERN BRAZIL Anthony W. Pereira
MANAGEMENT John Hendry MODERN CHINA Rana Mitter
NELSON MANDELA Elleke Boehmer MODERN DRAMA
MAO Delia Davin Kirsten E. Shepherd-Barr
MARINE BIOLOGY Philip V. Mladenov MODERN FRANCE
MARKETING Vanessa R. Schwartz
Kenneth Le Meunier-FitzHugh MODERN INDIA Craig Jeffrey
THE MARQUIS DE SADE John Phillips MODERN IRELAND Senia Pašeta
MARTYRDOM Jolyon Mitchell MODERN ITALY Anna Cento Bull
MARX Peter Singer MODERN JAPAN
MATERIALS Christopher Hall Christopher Goto-Jones
MATHEMATICAL ANALYSIS MODERN LATIN AMERICAN
Richard Earl LITERATURE
MATHEMATICAL FINANCE Roberto González Echevarría
Mark H. A. Davis MODERN WAR Richard English
MATHEMATICS Timothy Gowers MODERNISM Christopher Butler
MATTER Geoff Cottrell MOLECULAR BIOLOGY Aysha Divan
THE MAYA Matthew Restall and and Janice A. Royds
Amara Solari MOLECULES Philip Ball
THE MEANING OF LIFE MONASTICISM Stephen J. Davis
Terry Eagleton THE MONGOLS Morris Rossabi
MEASUREMENT David Hand MONTAIGNE William M. Hamlin
MEDICAL ETHICS Michael Dunn and MOONS David A. Rothery
Tony Hope MORMONISM Richard Lyman Bushman
MEDICAL LAW Charles Foster MOUNTAINS Martin F. Price
MEDIEVAL BRITAIN John Gillingham MUHAMMAD Jonathan A. C. Brown
and Ralph A. Griffiths MULTICULTURALISM Ali Rattansi
MEDIEVAL LITERATURE MULTILINGUALISM John C. Maher
Elaine Treharne MUSIC Nicholas Cook
MEDIEVAL PHILOSOPHY MUSIC AND TECHNOLOGY
John Marenbon Mark Katz
MEMORY Jonathan K. Foster MYTH Robert A. Segal
METAPHYSICS Stephen Mumford NANOTECHNOLOGY Philip Moriarty
METHODISM William J. Abraham NAPOLEON David A. Bell
THE MEXICAN REVOLUTION THE NAPOLEONIC WARS
Alan Knight Mike Rapport
MICROBIOLOGY Nicholas P. Money NATIONALISM Steven Grosby
MICROBIOMES Angela E. Douglas NATIVE AMERICAN LITERATURE
MICROECONOMICS Avinash Dixit Sean Teuton
MICROSCOPY Terence Allen NAVIGATION Jim Bennett
THE MIDDLE AGES Miri Rubin NAZI GERMANY Jane Caplan
MILITARY JUSTICE Eugene R. Fidell NEGOTIATION Carrie Menkel-Meadow
MILITARY STRATEGY NEOLIBERALISM Manfred B. Steger
Antulio J. Echevarria II and Ravi K. Roy
NETWORKS Guido Caldarelli and PENTECOSTALISM William K. Kay
Michele Catanzaro PERCEPTION Brian Rogers
THE NEW TESTAMENT THE PERIODIC TABLE Eric R. Scerri
Luke Timothy Johnson PHILOSOPHICAL METHOD
THE NEW TESTAMENT AS Timothy Williamson
LITERATURE Kyle Keefer PHILOSOPHY Edward Craig
NEWTON Robert Iliffe PHILOSOPHY IN THE ISLAMIC
NIETZSCHE Michael Tanner WORLD Peter Adamson
NINETEENTH‑CENTURY BRITAIN PHILOSOPHY OF BIOLOGY
Christopher Harvie and Samir Okasha
H. C. G. Matthew PHILOSOPHY OF LAW
THE NORMAN CONQUEST Raymond Wacks
George Garnett PHILOSOPHY OF MIND
NORTH AMERICAN INDIANS Barbara Gail Montero
Theda Perdue and Michael D. Green PHILOSOPHY OF PHYSICS
NORTHERN IRELAND David Wallace
Marc Mulholland PHILOSOPHY OF SCIENCE
NOTHING Frank Close Samir Okasha
NUCLEAR PHYSICS Frank Close PHILOSOPHY OF RELIGION
NUCLEAR POWER Maxwell Irvine Tim Bayne
NUCLEAR WEAPONS PHOTOGRAPHY Steve Edwards
Joseph M. Siracusa PHYSICAL CHEMISTRY Peter Atkins
NUMBER THEORY Robin Wilson PHYSICS Sidney Perkowitz
NUMBERS Peter M. Higgins PILGRIMAGE Ian Reader
NUTRITION David A. Bender PLAGUE Paul Slack
OBJECTIVITY Stephen Gaukroger PLANETARY SYSTEMS
OBSERVATIONAL ASTRONOMY Raymond T. Pierrehumbert
Geoff Cottrell PLANETS David A. Rothery
OCEANS Dorrik Stow PLANTS Timothy Walker
THE OLD TESTAMENT PLATE TECTONICS Peter Molnar
Michael D. Coogan PLATO Julia Annas
THE ORCHESTRA D. Kern Holoman POETRY Bernard O’Donoghue
ORGANIC CHEMISTRY POLITICAL PHILOSOPHY David Miller
Graham Patrick POLITICS Kenneth Minogue
ORGANIZATIONS Mary Jo Hatch POLYGAMY Sarah M. S. Pearsall
ORGANIZED CRIME POPULISM Cas Mudde and
Georgios A. Antonopoulos and Cristóbal Rovira Kaltwasser
Georgios Papanicolaou POSTCOLONIALISM
ORTHODOX CHRISTIANITY Robert J. C. Young
A. Edward Siecienski POSTMODERNISM Christopher Butler
OVID Llewelyn Morgan POSTSTRUCTURALISM
PAGANISM Owen Davies Catherine Belsey
PAKISTAN Pippa Virdee POVERTY Philip N. Jefferson
THE PALESTINIAN-ISRAELI PREHISTORY Chris Gosden
CONFLICT Martin Bunton PRESOCRATIC PHILOSOPHY
PANDEMICS Christian W. McMillen Catherine Osborne
PARTICLE PHYSICS Frank Close PRIVACY Raymond Wacks
PAUL E. P. Sanders PROBABILITY John Haigh
IVAN PAVLOV Daniel P. Todes PROGRESSIVISM Walter Nugent
PEACE Oliver P. Richmond PROHIBITION W. J. Rorabaugh
PROJECTS Andrew Davies RUSSELL A. C. Grayling
PROTESTANTISM Mark A. Noll THE RUSSIAN ECONOMY
PSEUDOSCIENCE Michael D. Gordin. Richard Connolly
PSYCHIATRY Tom Burns RUSSIAN HISTORY Geoffrey Hosking
PSYCHOANALYSIS Daniel Pick RUSSIAN LITERATURE Catriona Kelly
PSYCHOLOGY Gillian Butler and THE RUSSIAN REVOLUTION
Freda McManus S. A. Smith
PSYCHOLOGY OF MUSIC SAINTS Simon Yarrow
Elizabeth Hellmuth Margulis SAMURAI Michael Wert
PSYCHOPATHY Essi Viding SAVANNAS Peter A. Furley
PSYCHOTHERAPY Tom Burns and SCEPTICISM Duncan Pritchard
Eva Burns-Lundgren SCHIZOPHRENIA Chris Frith and
PUBLIC ADMINISTRATION Eve Johnstone
Stella Z. Theodoulou and Ravi K. Roy SCHOPENHAUER
PUBLIC HEALTH Virginia Berridge Christopher Janaway
PURITANISM Francis J. Bremer SCIENCE AND RELIGION
THE QUAKERS Pink Dandelion Thomas Dixon and Adam R. Shapiro
QUANTUM THEORY SCIENCE FICTION David Seed
John Polkinghorne THE SCIENTIFIC REVOLUTION
RACISM Ali Rattansi Lawrence M. Principe
RADIOACTIVITY Claudio Tuniz SCOTLAND Rab Houston
RASTAFARI Ennis B. Edmonds SECULARISM Andrew Copson
READING Belinda Jack SEXUAL SELECTION Marlene Zuk and
THE REAGAN REVOLUTION Gil Troy Leigh W. Simmons
REALITY Jan Westerhoff SEXUALITY Véronique Mottier
RECONSTRUCTION Allen C. Guelzo WILLIAM SHAKESPEARE
THE REFORMATION Peter Marshall Stanley Wells
REFUGEES Gil Loescher SHAKESPEARE’S COMEDIES
RELATIVITY Russell Stannard Bart van Es
RELIGION Thomas A. Tweed SHAKESPEARE’S SONNETS AND
RELIGION IN AMERICA Timothy Beal POEMS Jonathan F. S. Post
THE RENAISSANCE Jerry Brotton SHAKESPEARE’S TRAGEDIES
RENAISSANCE ART Stanley Wells
Geraldine A. Johnson GEORGE BERNARD SHAW
RENEWABLE ENERGY Nick Jelley Christopher Wixson
REPTILES T. S. Kemp MARY SHELLEY Charlotte Gordon
REVOLUTIONS Jack A. Goldstone THE SHORT STORY Andrew Kahn
RHETORIC Richard Toye SIKHISM Eleanor Nesbitt
RISK Baruch Fischhoff and John Kadvany SILENT FILM Donna Kornhaber
RITUAL Barry Stephenson THE SILK ROAD James A. Millward
RIVERS Nick Middleton SLANG Jonathon Green
ROBOTICS Alan Winfield SLEEP Steven W. Lockley and
ROCKS Jan Zalasiewicz Russell G. Foster
ROMAN BRITAIN Peter Salway SMELL Matthew Cobb
THE ROMAN EMPIRE ADAM SMITH Christopher J. Berry
Christopher Kelly SOCIAL AND CULTURAL
THE ROMAN REPUBLIC ANTHROPOLOGY
David M. Gwynn John Monaghan and Peter Just
ROMANTICISM Michael Ferber SOCIAL PSYCHOLOGY
ROUSSEAU Robert Wokler Richard J. Crisp
SOCIAL WORK Sally Holland and TRANSLATION Matthew Reynolds
Jonathan Scourfield THE TREATY OF VERSAILLES
SOCIALISM Michael Newman Michael S. Neiberg
SOCIOLINGUISTICS John Edwards TRIGONOMETRY
SOCIOLOGY Steve Bruce Glen Van Brummelen
SOCRATES C. C. W. Taylor THE TROJAN WAR Eric H. Cline
SOFT MATTER Tom McLeish TRUST Katherine Hawley
SOUND Mike Goldsmith THE TUDORS John Guy
SOUTHEAST ASIA James R. Rush TWENTIETH‑CENTURY BRITAIN
THE SOVIET UNION Stephen Lovell Kenneth O. Morgan
THE SPANISH CIVIL WAR TYPOGRAPHY Paul Luna
Helen Graham THE UNITED NATIONS
SPANISH LITERATURE Jo Labanyi Jussi M. Hanhimäki
THE SPARTANS Andrew J. Bayliss UNIVERSITIES AND COLLEGES
SPINOZA Roger Scruton David Palfreyman and Paul Temple
SPIRITUALITY Philip Sheldrake THE U.S. CIVIL WAR Louis P. Masur
SPORT Mike Cronin THE U.S. CONGRESS Donald A. Ritchie
STARS Andrew King THE U.S. CONSTITUTION
STATISTICS David J. Hand David J. Bodenhamer
STEM CELLS Jonathan Slack THE U.S. SUPREME COURT
STOICISM Brad Inwood Linda Greenhouse
STRUCTURAL ENGINEERING UTILITARIANISM
David Blockley Katarzyna de Lazari-Radek and
STUART BRITAIN John Morrill Peter Singer
SUBURBS Carl Abbott UTOPIANISM Lyman Tower Sargent
THE SUN Philip Judge VATICAN II Shaun Blanchard and
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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
by law, by licence or under terms agreed with the appropriate reprographics
rights organization. Enquiries concerning reproduction outside the scope of the
above should be sent to the Rights Department, Oxford University Press, at the
address above
You must not circulate this work in any other form
and you must impose this same condition on any acquirer
Published in the United States of America by Oxford University Press
198 Madison Avenue, New York, NY 10016, United States of America
British Library Cataloguing in Publication Data
Data available
Library of Congress Control Number: 2023930635
ISBN 978–0–19–285670–8
Printed and bound by
CPI Group (UK) Ltd, Croydon, CR0 4YY
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 From Jonathan M. W. Slack,


David Low’s The Breeds of Essential Developmental Biology,
3rd Edition, John Wiley & Sons
Domestic Animals of the British
Islands, London, 1842 3
9 The effect of gene drive
Heritage Images/Glowimages.com
on a population 39
Springer Nature
2 Mendel’s peas 6

10 Keto-enol 42
3 DNA two-­dimensional
structure 13
11 Romanov family 52
Heritage Images/Glowimages.com
4 DNA three-­dimensional
structure 19
12 Common ancestors 73
Creative Commons
Attribution-Share Alike 3.0
13 Human migrations out of
5 Protein synthesis 21 Africa as revealed by the
study of genetic markers
6 Gene structure 24 in populations 75

7 Behaviour of chromosomes 14 A Neanderthal face


on cell division 26 reconstructed by anatomical
From Jonathan M. W. Slack, methods and a Denisovan face
Essential Developmental Biology, reconstructed from the skull
3rd Edition, John Wiley & Sons structure deduced from the
DNA methylation pattern 85
8 Development 30 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
no related content on Scribd:
Le 16. — Émeute, sang, bruit de canons, bruit de mort. Nouvelle
venue comme un coup de foudre dans notre désert et calme
journée. Maurice, Caro, amis de Paris, je suis en peine, je vous vois
sur le volcan. Mon Dieu ! Je viens d’écrire à Caro et commence un
mot à M. d’Aurevilly, mon second frère en intérêt.

Le 18. — Point de lettre hier ni d’écriture ici. Je n’ai fait


qu’attendre, attendre un mécompte. Triste fin d’une journée
d’espérance, qui revient encore aujourd’hui ; rien ne peut l’éloigner
du cœur, cette trompeuse.
Je vais lire : que lirai-je ? Le choix des livres, malaisé comme
celui des hommes : peu de vrais et d’aimables.

Le 19. — Une lettre de Louise, pleine d’intérêt pour toi : rien que
cœur, esprit, charme d’un bout à l’autre, façon de dire qui ne se dit
nulle part que dans ces rochers de Rayssac. La solitude fait cela ; il
y vient des idées qui ne ressemblent à rien du monde, inconnues,
jolies comme des fleurs ou des mousses. Charmante Louise, que je
l’aime ! Je la trouve cette fois d’un calme, d’un désabusé qui
m’étonne, elle si illusionnée d’ordinaire. Je vais joindre l’autre
Louise, qui ressemble tant à celle-ci, ne trouves-tu pas ? et qui prie
aussi et fait prier pour ta guérison. « L’autre jour, m’écrit-elle (Louise
de Rayssac), j’étais à la Platée, paroisse de ma tante ; je
m’approchai d’une sainte fille qui habite cette église depuis le matin
jusqu’au soir, et qui est en grande vénération de sainteté. Je
soulevai un coin de son voile noir et lui dis bien bas : « Pardon,
mademoiselle, je voudrais vous demander des prières pour un jeune
homme malade, frère de la personne que j’aime le plus au monde. »
— « Eh bien ! je prierai », me dit-elle, avec cet air de modestie qui
donne encore plus de confiance à ma recommandation. — Je ne l’ai
pas revue. »
N’est-ce pas un joli trait pieux, mon ami, cette jeune fille quêtant
pour toi des prières avec un air d’intérêt céleste ? Elle est
charmante. Les anges lui auraient donné.

Le 21. — Mon bonheur, mon charme, mes délices, écrire au


soleil, écouter les oiseaux.
Ce n’a pas été long ce beau jour de ce matin. Hélas ! mon ami,
une lettre de Caro m’est venue parler si tristement de ta santé que
j’en suis accablée. Il tousse, il tousse encore ! Ces mots retentissent
partout depuis, une pensée désolante me poursuit, passe et repasse
dedans, dehors, et va tomber sur un cimetière ; je ne puis voir une
feuille verte sans penser qu’elle tombera bientôt et qu’alors les
poitrinaires meurent. Mon Dieu, détournez ces pressentiments,
guérissez-moi ce pauvre frère ! Que me faudrait-il faire pour lui ?
Impuissante affection ! Tout se réduit pour moi à souffrir pour toi.

Le 22. — Si jamais tu lis ceci, mon ami, tu auras l’idée d’une


affection permanente, ce quelque chose pour quelqu’un qui vous
occupe au coucher, au lever, dans le jour et toujours, qui fait
tristesse ou joie mobile et centre de l’âme. — En lisant un livre de
géologie, j’ai rencontré un éléphant fossile découvert dans la
Laponie, et une pirogue déterrée dans l’île des Cygnes, en creusant
les fondations du pont des Invalides. Me voilà sur l’éléphant, me
voilà dans la pirogue, faisant le tour des mers du Nord et de l’île des
Cygnes, voyant ces lieux du temps de ces choses : la Laponie
chaude, verdoyante et peuplée, non de nains, mais d’hommes
beaux et grands, de femmes s’en allant en promenade sur un
éléphant, dans ces forêts, sous ces monts pétrifiés aujourd’hui ; et
l’île des Cygnes, blanche de fleurs, et de leur duvet, oh ! que je la
trouve belle ! Et ses habitants, qui sont-ils ? que font-ils dans ce coin
du globe ? Descendants comme nous de l’exilé d’Éden, connaissent-
ils sa naissance, sa vie, sa chute, sa lamentable et merveilleuse
histoire ; cette Ève pour laquelle il a perdu le ciel, tant de malheur et
de bonheur ensemble, tant d’espérances dans la foi, tant de larmes
sur leurs enfants, tant et tant de choses que nous savons, que savait
peut-être avant nous ce peuple dont il ne reste qu’une planche ?
Naufrages de l’humanité que Dieu seul connaît, dont il a caché les
débris dans les profondeurs de la terre, comme pour les dérober à
notre curiosité ! S’il en laisse voir quelque chose, c’est pour nous
apprendre que ce globe est un abîme de malheurs, et que ce qu’on
gagne à remuer ses entrailles, c’est de découvrir des inscriptions
funéraires, des cimetières. La mort est au fond de tout, et on creuse
toujours comme qui cherche l’immortalité.
Une lettre de Félicité, qui ne m’apprend rien de meilleur de toi.
Quand écriront-ils, ceux qui en savent davantage ? Si on voyait
battre un cœur de femme, on en aurait plus de pitié. Pourquoi
sommes-nous ainsi, qu’un désir nous consume, qu’une crainte nous
brise, qu’une attente nous obsède, qu’une pensée nous remplisse et
que tout ce qui nous touche nous fasse tressaillir ? Souvenir de
lettres, heure de la poste, vue d’un papier, Dieu sait ce que j’en
éprouve ! Le désert des Coques aura vu bien des choses pour toi.
Ma douce amie, ma sœur de peines et d’affections est là, pour mon
bonheur, d’un côté, pour m’attrister, de l’autre, quand je la vois
souffrir, et qu’il me faut lui cacher mes souffrances pour ménager sa
sensibilité.

Le 24. — Inquiétudes, alarmes croissantes, lettre de M. de


Frégeville qui t’a trouvé plus mal. Mon Dieu ! faut-il apprendre
comme par hasard que je puis te perdre ? Personne de plus près
qu’un étranger ne me parlera pas de toi, ne me dira pas qu’il t’a vu
pour moi ! Dans l’éloignement, rien n’est accablant comme le
silence. C’est la mort avancée. Mon ami, mon frère, mon cher
Maurice, je ne sais que penser, que dire, que sentir. Après Dieu, je
ne vis qu’en toi comme une martyre, en souffrant. Et qu’est-ce que
cela, si je pouvais l’offrir pour te racheter ? quand je plongerais dans
une mer de douleur pour te sauver du naufrage. Toute rédemption
se fait par la souffrance : acceptez la mienne, mon Dieu, unissez-la
à celle des sœurs de Lazare, unissez-la à celle de Marie, au glaive
qui perça son âme auprès de Jésus mourant ; acceptez, mon Dieu,
coupez, tranchez en moi, mais qu’il se fasse une résurrection !
Le 25. — Courrier passé sans me rien laisser. Mêmes doutes et
incertitudes, mêmes craintes envahissantes. Savoir et ne pas
savoir ! État d’indicibles angoisses. Et voilà la fin de ce cahier : mon
Dieu ! qui le lira [29] ?
[29] Qui devait le lire ? Ainsi qu’Eugénie de Guérin le
pressentait, ce ne fut pas Maurice, qui, ramené par elle,
et non sans peine, au Cayla, s’y éteignit moins de deux
mois après la date de cette page, le 19 juillet 1839. On
trouvera dans un des cahiers qui suivent le touchant récit
des derniers instants d’un frère si tendrement aimé.
IX

ENCORE A LUI
A MAURICE MORT, A MAURICE AU CIEL

IL ÉTAIT LA GLOIRE ET LA JOIE DE MON CŒUR.

OH ! QUE C’EST UN DOUX NOM ET PLEIN DE DILECTION QUE LE NOM DE


FRÈRE !

Vendredi 19 juillet, à 11 heures ½, date éternelle !

Le 21 juillet [1839]. — Non, mon ami, la mort ne nous séparera


pas, ne t’ôtera pas de ma pensée : la mort ne sépare que le corps ;
l’âme, au lieu d’être là, est au ciel, et ce changement de demeure
n’ôte rien à ses affections. Bien loin de là, j’espère ; on aime mieux
au ciel où tout se divinise. O mon ami, Maurice, Maurice, es-tu loin
de moi, m’entends-tu ? Qu’est-ce que les lieux où tu es maintenant ?
qu’est-ce que Dieu si beau, si puissant, si bon, qui te rend heureux
par sa vue ineffable en te dévoilant l’éternité ? Tu vois ce que
j’attends, tu possèdes ce que j’espère, tu sais ce que je crois.
Mystères de l’autre vie, que vous êtes profonds, que vous êtes
terribles, que quelquefois vous êtes doux ! oui, bien doux, quand je
pense que le ciel est le lieu du bonheur. Pauvre ami, tu n’en as eu
guère ici-bas, de bonheur ; ta vie si courte n’a pas eu le temps du
repos. O Dieu ! soutenez-moi, établissez mon cœur dans la foi.
Hélas ! je n’ai pas assez de cet appui. Que nous t’avons gardé et
caressé et baisé, ta femme et nous tes sœurs, mort dans ton lit, la
tête appuyée sur un oreiller comme si tu dormais ! Puis nous t’avons
suivi dans le cimetière, dans la tombe, ton dernier lit, prié et pleuré,
et nous voici, moi t’écrivant comme dans une absence, comme
quand tu étais à Paris. Mon ami, est-il vrai, ne te reverrons-nous plus
nulle part sur la terre ? Oh ! moi je ne veux pas te quitter ; quelque
chose de doux de toi me fait présence, me calme, fait que je ne
pleure pas. Quelquefois larmes à torrents, puis l’âme sèche. Est-ce
que je ne le regretterais pas ? Toute ma vie sera de deuil, le cœur
veuf, sans intime union. J’aime beaucoup Marie et le frère qui me
reste, mais ce n’est pas avec notre sympathie. Reçu une lettre de
ton ami d’Aurevilly pour toi. Déchirante lettre arrivée sur ton cercueil.
Que cela m’a fait sentir ton absence ! Il faut que je quitte ceci, ma
tête n’y tient pas, parfois je me sens des ébranlements de cerveau.
Que n’ai-je des larmes ! J’y noierais tout.

Le 22. — Sainte Madeleine aujourd’hui, celle à qui il a été


beaucoup pardonné parce qu’elle a beaucoup aimé. Que cette
pensée, qui m’est venue pendant la messe que nous avons
entendue pour toi, m’a consolée sur ton âme ! Oh ! cette âme aura
été pardonnée, mon Dieu, je me souviens de tout un temps de foi et
d’amour qui n’aura pas été perdu devant vous.

Où l’éternité réside
On retrouve jusqu’au passé.

Le passé de la vertu surtout, qui doit couvrir les faiblesses, les


erreurs présentes. Oh ! que ce monde, cet autre monde où tu es
m’occupe. Mon ami, tu m’élèves en haut, mon âme se détache de
plus en plus de la terre ; la mort, je crois, me ferait plaisir.
— Eh ! que ferions-nous de l’éternité en ce monde ? Visites de
ma tante Fontenilles, d’Éliza, de M. Limer, d’Hippolyte, de Thérèse,
tout monde, hélas ! qui devait venir en joie de noces, et qui sont là
pour un enterrement. Ainsi changent les choses. Ainsi Dieu le veut.
Bonsoir, mon ami. Oh ! que nous avons prié ce matin sur ta tombe,
ta femme, ton père et tes sœurs !
Des visites, toujours des visites. Oh ! qu’il est triste de voir des
vivants, d’entrer en conversation, de revoir le cours ordinaire des
choses, quand tout est changé au cœur ! Mon pauvre ami, quel vide
tu me fais ! Partout ta place sans t’y voir… Ces jeunes filles, ces
jeunes gens, nos parents, nos voisins, qui remplissent en ce
moment le salon, qui sont autour de toi mort, t’entoureraient vivant et
joyeux, car tu te plaisais avec eux, et leur jeune gaieté t’égayait.
Lettre touchante de l’abbé de Rivières, qui te pleure en ami ;
pareille lettre de sa mère pour moi. Expression la plus tendre de
regret, douleur de mère mêlée à la mienne. Oh ! elle savait que tu
étais le fils de mon cœur.
Au retour de…

Sans date. — Je ne sais ce que j’allais dire hier à cet endroit


interrompu. Toujours larmes et regrets. Cela ne passe pas, au
contraire : les douleurs profondes sont comme la mer, avancent,
creusent toujours davantage. Huit soirs ce soir que tu reposes là-
bas, à Andillac, dans ton lit de terre. O Dieu, mon Dieu ! consolez-
moi ! Faites-moi voir et espérer au delà de la tombe, plus haut que
n’est tombé ce corps. Le ciel, le ciel ! oh ! que mon âme monte au
ciel !
Aujourd’hui grande venue de lettres que je n’ai pas lues. Que lire
là dedans ? Des mots qui ne disent rien. Toute consolation humaine
est vide. Que j’éprouve cruellement la vérité de ces paroles de
l’Imitation ! Ta berceuse est venue, la pauvre femme, toute larmes, et
portant gâteaux et figues que tu aurais mangés. Quel chagrin m’ont
donné ces figues ! Le plus petit plaisir que je te vois venir me semble
immense. Et le ciel si beau, et les cigales, le bruit des champs, la
cadence des fléaux sur l’aire, tout cela qui te charmerait me désole.
Dans tout je vois la mort. Cette femme, cette berceuse qui t’a veillé
et tenu un an malade sur ses genoux, m’a porté plus de douleur que
n’eût fait un drap mortuaire. Déchirante apparition du passé :
berceau et tombe. Je passerais la nuit ici avec toi sur ce papier ;
mais l’âme veut prier, l’âme te fera plus de bien que le cœur.
Chaque fois que je pose la plume ici, une lame me passe au
cœur. Je ne sais si je continuerai d’écrire. A quoi sert ce Journal ?
Pour qui ? hélas ! Et cependant je l’aime, comme on aime une boîte
funèbre, un reliquaire où se trouve un cœur mort, tout embaumé de
sainteté et d’amour. Ainsi ce papier où je te conserve, ami tant aimé,
où je te garde un parlant souvenir, où je te retrouverai dans ma
vieillesse… si je vieillis. Oh oui ! viendront les jours où je n’aurai de
vie que dans le passé, le passé avec toi, près de toi jeune,
intelligent, aimable, sensibilisant tout ce qui t’approchait, tel que je te
vois, tel que tu nous as quittés. Maintenant je ne sais ce qu’est ma
vie, si je vis. Tout est changé au dedans, au dehors. O mon Dieu !
que ces lettres sont déchirantes, ces lettres du bon marquis et de
ton ami surtout. Oh ! celles-ci, qu’elles m’ont fait pleurer ! Il y a là
dedans tant de larmes pour mes larmes ! Cet intime ami me touche
comme ferait te voir. Mon cher Maurice, tout ce que tu as aimé m’est
cher, me semble une portion de toi-même. Frère et sœur nous
serons avec M. d’Aurevilly ; il se dit mon frère.
Lu les Confessions de saint Augustin à l’endroit de la mort de son
ami. Trouvé un charme de vérité, une saillante expression de
douleur à cette lecture qui m’a fait du bien. Les saints savent
toujours mêler quelque chose de consolant à leurs larmes.

Le 28. — Rien n’est poignant comme le retour des mêmes


personnes dans des jours tout différents, revoir en deuil qui vous
avait porté la joie. Sa tante, la tante de Caroline, celle qui, il y a deux
ans, nous amenait ta fiancée, est arrivée, est ici où tu n’es pas…

Le 4 août. — A pareil jour vint au monde un frère que je devais


bien aimer, bien pleurer, hélas ! ce qui va souvent ensemble. J’ai vu
son cercueil dans la même chambre, à la même place où, toute
petite, je me souviens d’avoir vu son berceau, quand on m’amena de
Gaillac où j’étais, pour son baptême. Ce baptême fut pompeux, plein
de fête, plus qu’aucun autre de nous, marqué de distinction. Je jouai
beaucoup et je repartis le lendemain, aimant fort ce petit enfant qui
venait de naître. J’avais cinq ans. Deux ans après je revins, lui
portant une robe que je lui avais faite. Je lui mis sa robe et le menai
par la main le long de la garenne du nord, où il fit quelques pas tout
seul, les premiers, ce que j’allai annoncer en grande joie à ma
mère : « Maurice, Maurice a marché seul ! » Souvenir qui me vient
tout mouillé de larmes.

Le 6. — Journée de prières et de pieuse consolation : pèlerinage


de ton ami, le saint abbé de Rivières, à Andillac, où il a dit la messe,
où il est venu prier avec tes sœurs près de ta tombe. Oh ! que cela
m’a touchée ; que j’ai béni dans mon cœur ce pieux ami agenouillé
sur tes restes, dont l’âme, par delà ce monde, soulageait la tienne
souffrante, si elle souffre ! Maurice, je te crois au ciel. Oh ! j’ai cette
confiance, que tes sentiments religieux me donnent, que la
miséricorde de Dieu m’inspire. Dieu si bon, si compatissant, si
aimant, si Père, n’aurait-il pas eu pitié et tendresse pour un fils
revenu à lui ? Oh ! il y a trois ans qui m’affligent ; je voudrais les
effacer de mes larmes. Mon Dieu, tant de supplications ont été
faites ! Mon Dieu, vous les avez entendues, vous les aurez
exaucées. O mon âme, pourquoi es-tu triste et pourquoi me
troubles-tu ?

Le 13. — Besoin d’écrire, besoin de penser, besoin d’être seule,


non pas seule, avec Dieu et toi. Je me trouve isolée au milieu de
tous. O solitude vivante, que tu seras longue !

Le 17. — Commencé à lire les Saints désirs de la mort, lecture


de mon goût. Mon âme vit dans un cercueil. Oh ! oui, enterrée,
ensevelie en toi, mon ami ; de même que je vivais en ta vie, je suis
morte en ta mort. Morte à tout bonheur, à toute espérance ici-bas.
J’avais tout mis en toi, comme une mère en son fils ; j’étais moins
sœur que mère. Te souviens-tu que je me comparais à Monique
pleurant son Augustin, quand nous parlions de mes afflictions pour
ton âme, cette chère âme dans l’erreur ? Que j’ai demandé à Dieu
son salut, prié, supplié ! Un saint prêtre me dit : « Votre frère
reviendra. » Oh ! il est revenu, et puis m’a quittée pour le ciel, pour le
ciel, j’espère. Il y a eu des signes évidents de grâce, de miséricorde
dans cette mort. Mon Dieu, j’ai plus à vous bénir qu’à me plaindre.
Vous en avez fait un élu par les souffrances qui rachètent, par
l’acceptation et résignation qui méritent, par la foi qui sanctifie. Oh !
oui, cette foi lui était revenue vive et profonde ; cela s’est vu dans
des actes religieux, des prières, des lectures, et dans ce baiser à la
croix fait avec tant d’âme et d’amour un peu avant de mourir ! Oh !
moi qui le voyais faire, qui le regardais tant dans ses dernières
actions, j’ai dit, mon Dieu, j’ai dit qu’il s’en allait en paradis. Ainsi
finissent ceux qui s’en vont dans la vie meilleure.
Maurice, mon ami, qu’est-ce que le ciel, ce lieu des amis ?
Jamais ne me donneras-tu signe de là ? Ne t’entendrai-je pas,
comme on dit que quelquefois on entend les morts ? Oh ! si tu le
pouvais, s’il existe quelque communication entre ce monde et l’autre,
reviens ! Je n’aurai pas peur un soir de voir une apparition, quelque
chose de toi à moi qui étions si unis. Toi au ciel et moi sur la terre,
oh ! que la mort nous sépare ! J’écris ceci à la chambrette, cette
chambrette tant aimée où nous avons tant causé ensemble, rien que
nous deux. Voilà ta place et là la mienne. Ici était ton portefeuille si
plein de secrets de cœur et d’intelligence, si plein de toi et de choses
qui ont décidé de ta vie. Je le crois, je crois que les événements ont
influé sur ton existence. Si tu étais demeuré ici, tu ne serais pas
mort. Mort ! terrible et unique pensée de ta sœur.

Le 20. — Hier allée à Cahuzac entendre la messe pour toi en


union de celle que le prince de Hohenlohe offrait en Allemagne pour
demander à Dieu ta guérison, hélas ! demandée trop tard. Quinze
jours après ta mort, la réponse est venue m’apporter douleurs au
lieu d’espérance. Que de regrets de n’avoir pas pensé plus tôt à ce
moyen de salut, qui en a sauvé tant d’autres ! C’est sur des faits bien
établis que j’avais eu recours au saint thaumaturge, et je croyais tant
au miracle ! Mon Dieu, j’y crois encore, j’y crois en pleurant. Maurice,
un torrent de tristesse m’a passé sur l’âme aujourd’hui. Chaque jour
agrandit ta perte, agrandit mon cœur pour les regrets. Seule dans le
bois avec mon père, nous nous sommes assis à l’ombre, parlant de
toi. Je regardais l’endroit où tu vins t’asseoir il y a deux ans, le
premier jour, je crois, où tu fis quelques pas dehors. Oh ! quel
souvenir de maladie et de guérison ! Je suis triste à la mort. Je
voudrais te voir. Je prie Dieu à tout moment de me faire cette grâce.
Ce ciel, ce ciel des âmes, est-il si loin de nous, le ciel du temps de
celui de l’éternité ? O profondeur ! ô mystères de l’autre vie qui nous
sépare ! Moi qui étais si en peine sur lui, qui cherchais tant à tout
savoir, où qu’il soit maintenant, c’est fini. Je le suis dans les trois
demeures, je m’arrête aux délices, je passe aux souffrances, aux
gouffres de feu. Mon Dieu, mon Dieu, non ! Que mon frère ne soit
pas là, qu’il n’y soit pas ! Il n’y est pas ; son âme, l’âme de Maurice
parmi les réprouvés… Horrible crainte, non ! Mais au purgatoire où
l’on souffre, où s’expient les faiblesses du cœur, les doutes de l’âme,
les demi-volontés au mal. Peut-être mon frère est là qui souffre et
nous appelle dans les gémissements comme il faisait dans les
souffrances du corps : « Soulagez-moi, vous qui m’aimez. » Oui,
mon ami, par la prière. Je vais prier ; je l’ai tant fait et le ferai
toujours. Des prières, oh ! des prières pour les morts, c’est la rosée
du purgatoire.
Sophie m’a écrit, cette Sophie, amie de Marie, qui m’aime en elle
et vient me consoler. Mais rien d’humain ne console. Je voudrais
aller en Afrique porter ma vie à quelqu’un, m’employer au salut des
Arabes dans l’établissement de Mme Vialar. Mes jours ne me
sembleraient pas vides, inutiles comme ils sont. Cette idée de cloître
qui s’en était allée, qui s’était retirée devant toi, me revient.
Le rosier, le petit rosier des Coques, a fleuri. Que de tristesses,
de craintes, de souvenirs épanouis avec ces fleurs, renfermés dans
ce vase donné par Marie, emporté dans notre voyage, avec nous
dans la voiture de Tours à Bordeaux, de là ici ! Ce rosier te faisait
plaisir ; tu te plaisais à le voir, à penser d’où il venait. Je voyais cela
et comme étaient jolis ces petits boutons et cette petite verdure.

Le 22. — Mis au doigt la bague antique que tu avais prise et mise


ici il y a deux ans, cette bague qui nous avait tant de fois fait rire
quand je te disais : « Et la bague ? » Oh ! qu’elle m’est triste à voir et
que je l’aime ! Mon ami, tout m’est relique de toi.
La mort nous revêtira de toute chose. Consolante parole que je
viens de méditer, qui me revêt le cœur d’espérance, ce pauvre cœur
dépouillé.
Comme j’aime ses lettres, ces lettres qui ne viennent pas ! Mon
Dieu, recevez ce que j’en souffre et toutes les douleurs de cette
affection. Voilà que cette âme m’attriste, que son salut m’inquiète,
que je souffrirais le martyre pour lui mériter le ciel. Exaucez, mon
Dieu, mes prières : éclairez, attirez, touchez cette âme si faite pour
vous connaître et vous servir ! Oh ! quelle douleur de voir s’égarer de
si belles intelligences, de si nobles créatures, des êtres formés avec
tant de faveur, où Dieu semble avoir mis toutes ses complaisances
comme en des fils bien-aimés, les mieux faits à son image ! Ah !
qu’ils sont à plaindre ! que mon âme souvent les pleure avec Jésus
venu pour les sauver ! Je voudrais le salut de tous, que tous profitent
de la rédemption qui s’étend à tout le genre humain. Mais le cœur a
ses élus, et pour ceux-là on a cent fois plus de désirs et de crainte.
Cela n’est pas défendu. Jésus, n’aviez-vous pas votre Jean bien-
aimé, dont les apôtres disaient que, par amour, vous feriez qu’il ne
mourrait pas ? Faites qu’ils vivent toujours, ceux que j’aime, qu’ils
vivent de la vie éternelle ! Oh ! c’est pour cela, pas pour ici que je les
aime. A peine, hélas ! si l’on s’y voit. Je n’ai fait que l’apercevoir ;
mais l’âme reste dans l’âme.
Le 25. — Tristesse et communion ; pleuré en Dieu ; écrit à ton
ami ; lu Pascal, l’étonnant penseur. J’ai recueilli cette pensée sur
l’amour de Dieu, qu’on aime sans le connaître : Le cœur a ses
raisons que la raison ne comprend pas. Bien souvent j’ai senti cela.

Le 26. — Quelques gouttes de pluie sur la terre ardente. Peut-


être orage ce soir, ramassé par ces vapeurs. Qu’il tonne, qu’il passe
des torrents d’eau et de vent ! je voudrais du bruit, des secousses,
tout ce qui n’est pas ce calme affaissant. — Si j’écrivais sa vie, cette
vie si jeune, si riche, si rare, si rattachée à tant d’événements, à tant
d’intérêts, à tant de cœurs ! peu de vies semblables.

Le 27. — Je ne sais, sans mon père, j’irais peut-être joindre les


sœurs de Saint-Joseph, à Alger. Au moins ma vie serait utile. Qu’en
faire à présent ? Je l’avais mise en toi, pauvre frère ! Tu me disais de
ne pas te quitter. En effet, je suis bien demeurée près de toi pour te
voir mourir. Un ecce homo, l’homme de douleur, tous les autres
derrière celui-là. Souffrances de Jésus, saints désirs de la mort,
uniques pensées et méditations. Écrit à Louise comme à Marie ; il
fait bon écrire à celle-là. Et lui, pourquoi ne pas écrire, ton frère ?
Serait-il mort aussi ? Mon Dieu, que le silence m’effraye à présent :
pardonnez-moi tout ce qui me fait peur. L’âme qui vous est unie,
qu’a-t-elle à craindre ? Ne vous aimerais-je pas, mon Dieu, unique et
véritable et éternel amour ? Il me semble que je vous aime, comme
disait le timide Pierre, mais pas comme Jean, qui s’endormait sur
votre cœur. Divin repos qui me manque ! Que vais-je chercher dans
les créatures ? Me faire un oreiller d’une poitrine humaine, hélas ! j’ai
vu comme la mort nous l’ôte. Plutôt m’appuyer, Jésus, sur votre
couronne d’épines.

Le 28. — Saint Augustin aujourd’hui, ce saint qui pleurait si


tendrement son ami et d’avoir aimé Dieu si tard. Que je n’aie pas
ces deux regrets : oh ! que je n’aie pas cette douleur à deux
tranchants, qui me fendrait l’âme à la mort ! Mourir sans amour, c’est
mourir en enfer. Amour divin, seul véritable. Les autres ne sont que
des ombres.
Accablement, poids de douleurs ; essayons de soulever ce mont
de tristesse. Que faire ? Oh, que l’âme est ignorante ! Il faut
s’attacher à Dieu, à celui qui soulève et le vaisseau et la mer. Pauvre
nacelle, que je suis sur un océan de larmes !
Recueillir chaque jour une pensée. Voici celle d’aujourd’hui :
« C’est une chose horrible de sentir continuellement s’écouler ce
qu’on possède et qu’on puisse s’y attacher, sans avoir envie de
chercher s’il n’y a point quelque chose de permanent. » — Beaucoup
lu, soigné de petits oiseaux qu’on a apportés, sans goût, par pitié,
toutes mes affections mortes ; toutes, hormis celle que la mort m’a
prise.

Le 29. — L’homme est un roseau pensant.

Le 30. — Qu’il faisait bon ce matin dans la vigne, cette vigne aux
raisins-chasselas que tu aimais ! En m’y voyant, en mettant le pied
où tu l’avais mis, la tristesse m’a rempli l’âme. Je me suis assise à
l’ombre d’un cerisier, et là, pensant au passé, j’ai pleuré. Tout était
vert, frais, doré de soleil, admirable à voir. Ces approches d’automne
sont belles, la température adoucie, le ciel plus nuagé, des teintes
de deuil qui commencent. Tout cela, je l’aime, je m’en savoure l’œil,
m’en pénètre jusqu’au cœur, qui tourne aux larmes. Vu seule, c’est
si triste ! Toi, tu vois le ciel ! Oh ! je ne te plains pas. L’âme doit
goûter d’ineffables ravissements,

Se plongeant dans l’extase où fut l’aveugle-né


Quand le jour apparut à son œil étonné.
Le 31. — Quelle différence de ce que je dis à ce que je dirais s’il
vivait ! Mon Dieu, tout est changé en moi et hors de moi : la mort
étend quelque chose de noir sur toutes choses. — Écrit à Misy sur la
mort de son oncle Jules de Roquefeuil, disparu tout jeune de ce
monde. De tous côtés, des tombes s’ouvrent.
« Cet étrange secret dans lequel Dieu s’est retiré, impénétrable à
la vue de l’homme, est une grande leçon pour nous porter à la
solitude loin de la vue des hommes. »
« L’homme est ainsi fait qu’à force de lui dire qu’il est un sot, il le
croit ; et à force de se le dire à soi-même, on le croit… »
« … Dieu a créé l’homme avec deux amours : l’un pour Dieu,
l’autre pour soi-même… Le péché étant arrivé, l’homme a perdu le
premier de ces amours ; et l’amour pour soi-même étant resté seul
dans cette grande âme capable d’un amour infini, cet amour-propre
s’est étendu et débordé dans le vide que l’amour de Dieu a
laissé [30] . »
[30] Pascal, Pensées ; Lettre sur la mort de son père.

Il pleut ; cette pluie, qui reverdit prés et bois, tombe sur la terre
qui te couvre et dissout tes restes au cimetière, là-bas, à Andillac.
Qu’on est heureux de penser qu’il y a dans l’homme quelque chose
que n’atteint pas la destruction !
« Il est des créatures que vous retirez de ce monde pour de
légères faiblesses ; c’est par amour et pour leur sauver de nouvelles
chutes. » — Si on ne savait que cette pensée est de Shakspeare, on
la croirait de Fénelon. Oh ! je sais à qui je l’applique.

Le 5 septembre. — Une lettre de Marie, la triste Marie, qui récite


tous les jours l’office des morts. Ainsi le cœur de la femme : même
en se tournant vers Dieu, il regarde ses affections.
Le 9. — Le découragement me prend pour tout dans la vie. Je ne
continuerai pas d’écrire. A quoi bon ce mémorandum ? Pourquoi ?
puisque ce ne peut être pour lui ! Quand il vivait, j’avais en lui mon
soutien ; j’avais mon plaisir dans la pensée de lui faire plaisir. —
Cela ôté, que reste-t-il à ces distractions humaines, lectures,
pensées, poésie ? rien que leur valeur, qui n’est rien.
Écrit à Marie, autre poésie vivante encore. Je lui dis : « Croyez
que vous êtes aimée du cœur le plus mort. »

Le 25. — Encore à Marie.

Le 30. — A mon frère de Paris, le frère de celui de la tombe.


Plus d’écriture ici, plus de pensées ; l’illusion n’est plus possible ;
à chaque mot, à chaque ligne, je vois qu’il ne me lira pas. Mon Dieu,
j’avais tant l’habitude de lui tout dire ; je l’aimais tant ! « Le plus grand
malheur de la vie, c’est d’en rompre les relations. » Oh ! que
j’éprouve la vérité de ces mots, qui m’avaient frappée dans un livre
aux Coques.
J’ai besoin du Ciel.
Ce n’est pas pour rien que nous nous serons rencontrés dans la
vie. Je tâcherai, mon Dieu, de les tourner vers vous.

Je voudrais que le ciel fût tout tendu de noir,


Et qu’un bois de cyprès vînt à couvrir la terre ;
Que le jour ne fût plus qu’un soir.

Une gazelle errante


S’abrite en cette tour,
Et l’hirondelle y chante,
Y chante nuit et jour.
Le 3 octobre. — Écrit à Paris. Oh ! quel jour anniversaire de mon
départ l’an dernier ! — Dirai-je ici tous les souvenirs qui me viennent,
larmes, regrets, passé perdu, sitôt changé en deuil ? — Mon cœur
est plein, il veut pleurer. — Maurice, Maurice, n’est-ce pas vrai, les
pressentiments ? Quand je pense à ceux qui me tourmentaient dans
la route et à Paris et le jour de la noce, et qui se sont accomplis ! Je
rêvais mort ; je ne voyais que draperies mortuaires dans ce Salon où
l’on dansait, où je dansais dans ma tristesse, car je voulais écarter
ces pensées.
N’est-ce pas temps perdu que de rappeler ces choses, mon
Dieu ! Je suis seule devant vous : je pourrais mieux faire que de
m’affliger. N’êtes-vous pas là pour mon espérance, pour ma
consolation, pour me faire voir un monde meilleur où est mon frère ?

Le 4. — Je voulais envoyer à son ami deux grenades du


grenadier dont il a travaillé le pied quelques jours avant sa mort. Ce
fut son dernier mouvement sur la terre.

Le 6. — A l’heure qu’il est, midi, premier dimanche d’octobre,


j’étais à Paris, j’étais dans ses bras, place Notre-Dame-des-
Victoires. Un an passé, mon Dieu ! — Que je fus frappée de sa
maigreur, de sa toux, moi qui l’avais rêvé mort dans la route ! —
Nous allâmes ensemble à Saint-Sulpice à la messe à une heure.
Aujourd’hui à Lentin, dans la pluie, les poignants souvenirs et la
solitude… Mais, mon âme, apaise-toi avec ton Dieu que tu as reçu
dans cette petite église. C’est ton frère, ton ami, le bien-aimé
souverain que tu ne verras pas mourir, qui ne te manquera jamais ni
en cette vie ni en l’autre. Consolons-nous dans cette espérance, et
qu’en Dieu on retrouve tout ce qu’on a perdu. Si je pouvais m’en
aller en haut ; si je trouvais dans ma poitrine ce souffle qui vient le
dernier, ce souffle des mourants qui porte l’âme au ciel, oh ! je
n’aurais pas beaucoup de regrets à la vie. Mais la vie c’est une
épreuve, et la mienne est-elle assez longue ; ai-je assez souffert ?
Quand on se porte au Calvaire, on voit ce que coûte le ciel. Oh ! bien
des larmes, des déchirements, des épines, du fiel et du vinaigre. Ai-
je goûté de tout cela ? Mon Dieu, ôtez-moi la plainte, soutenez-moi
dans le silence et la résignation au pied de la Croix, avec Marie et
les femmes qui vous aimèrent.

Le 19. — Trois mois aujourd’hui de cette mort, de cette


séparation. Oh ! la douloureuse date, que néanmoins je veux écrire
chaque fois qu’elle reviendra. Il y a pour moi une si attachante
tristesse dans ce retour du 19, que je ne puis le voir sans le marquer
dans ma vie, puisque je note ma vie. Eh ! qu’y mettrais-je
maintenant, si je n’y mettais mes larmes, mes souvenirs, mes
regrets de ce que j’ai le plus aimé ? C’est tout ce qui vous viendra, ô
vous qui voulez que je continue ces cahiers, mon tous les jours au
Cayla. J’allais cesser de le faire, il y avait trop d’amertume à lui
parler dans la tombe ; mais puisque vous êtes là, frère vivant, et
avez plaisir de m’entendre, je continue ma causerie intime ; je
rattache à vous ce qui restait là, tombé brisé par la mort. J’écrirai
pour vous comme j’écrivais pour lui. Vous êtes mon frère d’adoption,
mon frère de cœur. Il y a là-dedans illusion et réalité, consolation et
tristesse : Maurice partout. C’est donc aujourd’hui 19 octobre que je
date pour vous et que je marque ce jour comme une époque dans
ma vie, ma vie d’isolement, de solitude, d’inconnue qui s’en va vers
quelqu’un du monde, vers vous à Paris, comme à peu près, je vous
l’ai dit, je crois, si Eustoquie, de son désert de Bethléem, eût écrit à
quelque élégant chevalier romain. Le contraste est piquant, mais ne
m’étonne pas. Quelqu’un, une femme, me disait qu’à ma place elle
serait bien embarrassée pour vous écrire. Moi, je ne comprends pas
pourquoi je le serais. Rien ne me gêne avec vous. En vérité, pas
plus qu’avec Maurice, vous m’êtes lui au cœur et à l’intelligence.
C’est à ce point de vue que se met notre intimité.

Le 20. — La belle matinée d’automne ! Un air transparent, un


lever du jour radieusement calme, des nuages en monceaux, du
nord au midi, des nuages d’un éclat, d’une couleur molle et vive, du
coton d’or sur un ciel bleu. C’était beau, c’était beau ! Je regrettais
d’être seule à le voir. J’ai pensé à notre peintre et ami, M. Augier, lui
qui sent si bien et prend sitôt le beau dans son âme d’artiste. Et puis
Maurice et puis vous, je vous aurais voulu voir tous sous mon ciel du
Cayla ; mais devons-nous nous rencontrer jamais plus sur la terre !
En allant au Pausadou, j’ai voulu prendre une fleur très-jolie. Je
l’ai laissée pour le retour, et j’ai passé par un autre chemin. Adieu
ma fleur. Quand j’y reviendrais, où serait-elle ? Une autre fois je ne
laisserai pas mes fleurs en chemin. Que de fois cependant cela
n’arrive-t-il pas dans la vie ?
Dimanche aujourd’hui. Revu à Andillac cette tombe toute
verdoyante d’herbe. Comme c’est venu vite, ces plantes ! Comme la
vie se hâte sur la mort, et que c’est triste à notre vue ! Que ce serait
désolant, sans la foi qui nous dit que nous devons renaître, sortir de
ces cimetières où nous semblons disparus !

Le 21. — Tonnerre, orage, tempête au dehors, mais calme au


dedans, ce calme d’une mer morte, qui a sa souffrance aussi bien
que l’agitation. Le repos n’est bon qu’en Dieu, ce repos des âmes
saintes qui, avant la mort, sont sorties de la vie. Heureux
dégagement ! Je meurs d’envie de tout ce qui est céleste : c’est
qu’ici-bas tout est vil et porte un poids de terre.
Lu quelques pages d’un voyage en Espagne. Singulier peuple de
brigands et de moines. Les moines sont tombés, que reste-t-il
maintenant ? Nous le voyons, des égorgeurs, Don Carlos à Bourges,
l’héritier de Ferdinand le Catholique mis hors du trône et du
royaume, prisonnier en France. Cette lecture m’intéresse. C’est
l’élégant journal d’un voyageur aimable, qui cause en courant, et
peint, avec le bon ton et la grâce d’esprit d’un homme du monde,
tout ce qu’il rencontre. Les lourdes descriptions m’assomment.
J’aime aussi M. de Custine, qui m’amuse, quoiqu’il soit parfois un
peu long ; mais c’est comme la longueur d’un bal. Puis il vient si peu
de livres au Cayla, que, pour peu qu’ils puissent plaire, ils plaisent
beaucoup.
Le 22. — Une lettre de Marie, de Marie ma sœur, qui m’a quittée
pour quelques jours avec Érembert. Me voici seule avec mon père.
Que notre famille est réduite, et je tremble en pensant que le cercle
peut encore se rétrécir !
Lu quelques passages des Saints Désirs de la Mort, livre
pieusement spirituel que j’aime, lecture qui porte au ciel. J’en ai
besoin pour mon âme qui tombe, qui s’affaisse sous le poids de la
vie. On peut se distraire dans le monde, mais les choses seules de
la foi soutiennent. Que je plains les âmes tristes qui ne savent pas
cela, ou ne le veulent pas croire ! J’en ai tant parlé à Maurice ; j’en
parle à tout ce que j’aime, des choses de l’éternité ; car, voyez-vous,
je n’aime pas pour ce monde, ce n’est pas la peine : c’est le ciel le
lieu de l’amour.

Le 24. — Lecture, ni écriture, ni prière ne peuvent empêcher les


larmes aujourd’hui. Mon pauvre Maurice ! Je me suis mise à penser
à tout ce qu’il a souffert, physiquement et dans l’âme, les derniers
temps de sa vie. Que cette vue est déchirante ! Mon Dieu, ne
l’aurez-vous pas soutenu ?

[Le 27.] — Nulle envie d’écrire depuis deux jours. Si je reprends


la plume aujourd’hui, c’est qu’en ouvrant mon portefeuille vert, j’ai vu
ce cahier et j’y mets que mon père vient de me remettre un paquet
de lettres de son cher Maurice, et de ses cheveux, pour les
renfermer, ces précieux restes, avec les autres que j’ai. O
enterrement ! Écrirai-je ce que je sens, ce que je pense, ce que je
souffre ? Je n’écris pas : je ne parlerais que du ciel et d’une tombe,
de ces choses qui ne doivent se dire qu’à Dieu.

Le 1er novembre. — Quel anniversaire ! J’étais à Paris, assise


seule dans le salon devant une table, pensant, comme à présent, à
cette fête des Saints. Il vint, Maurice, me trouver, causer un peu

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