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Bacteria: A Very Short Introduction
 VERY SHORT INTRODUCTIONS are for anyone wanting a stimulating and
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Very Short Introductions available now:

ABOLITIONISM Richard S. Newman

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

BACTERIA
A Very Short Introduction
SECOND EDITION
 Great Clarendon Street, Oxford, OX2 6DP, United Kingdom
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For Hilary
Contents

Preface

Acknowledgements

List of illustrations

1 Origins

2 Evolution

3 Microbiota and microbiome in humans

4 Discovery

5 Environment and civilization

6 Bacterial pathogenesis

7 Antibiotics

8 Antibiotic resistance

9 The future

Further reading
Index
Preface

It would be understandable if we thought that humans were the

principal species on this planet and that we now live in the era
where mammals dominate. As we consider previous ages, ending
with the Cretaceous extinction 65.5 million years ago, we could well
be forgiven for thinking that this was the ‘Age of the Dinosaurs’. The
reason is that we tend to classify each era with what can easily be
seen around us or from what palaeontologists have reported and
placed in museums of natural history for us to marvel at. The truth is
there never have been any dominant organisms other than bacteria
and that this planet has been in the ‘Age of Bacteria’ almost from the
very beginning when life emerged. Bacteria are the most numerous
of all organisms and their biomass is by far the largest on our planet
and has been estimated to be greater than all the rest combined.
Even within our own bodies the number of bacterial cells
outnumbers our own cells. They can survive almost everywhere on
the planet, from the coldest to the hottest places on earth, even to
the bottom of the oceans. No other organisms are as adaptable.
Acknowledgements

I should like to thank Dr H.-K. Young and Dr A. K. B. Rolfe for their

very useful scientific and medical advice and for their valuable
comments on the manuscript.
List of illustrations

1 Timescale of bacteria emergence

2 Structure of a T4 bacteriophage

3 Microbiome of the human body

4 Electron micrograph of a Gram-negative bacterium

Photo: Janice Haney Carr. CDC/National Escherichia, Shigella, Vibrio Reference Unit at
CDC

5 Pasteur’s swan-neck flask

6 Cheese production in the 14th century; illuminated page from the Tacuinum
Sanitas
Alinari via Getty Images

7 Basic flow chart of modern yogurt production

8 Diagram of a septic tank

9 Diagram of some of the pathogenicity factors associated with bacteria

Reproduced from Postgraduate Medical Journal, Wilson, J. W. et al., 78(918): 216–24,
2002, with permission from BMJ Publishing Group Ltd

10 Phagocytosis of bacterial cells by a macrophage

11 General comparison of endotoxin and exotoxin action

12 Engraving showing the masks used to prevent contracting plague in 1665
Courtesy of the Thomas Fisher Rare Book Library, University of Toronto

13 Graph showing the decline of bacterial infections over the last century

14 A chest X-ray of a person with pulmonary tuberculosis

Du Cane Medical Imaging Ltd/Science Photo Library

15 Sir Alexander Fleming in his laboratory at St Mary’s Hospital, London

Popperfoto/Getty Images

16 Diagram showing the stages of the chemical modification of penicillin

17 Quorum sensing

18 Diagram showing the structure of a biofilm

19 Mutant Prevention Concentration

Data from: Zhao and Drlica (2001) Clin. Infect Dis., 33: 147

20 Major mechanisms of resistance used by bacteria to overcome antibiotics

21 Vancomycin resistance in Enterococcus species

Data from Arthur and Courvalin (1993) Antimicrob Agents Chemother 37: 1563

22 Potential source of the vancomycin resistance vanA genes

Data from Marshall et al. (1998) Antimicrob Agents Chemother 42: 2215

23 eBURST diagram of the Staphylococcus aureus genomic sequences

Kindly provided by Prof. Ed Feil, University of Bath

24 Diagram showing the interactions between resistance in animals and in

humans
Phillips, I., Casewell, M., Cox, T., et al. (2004) Does the use of antibiotics in food
animals pose a risk to human health? A critical review of published data, Journal of
Antimicrobial Chemotherapy, 53, 28–52. With permission from Oxford University
Press

25 Micrograph of the synthetic bacterium Mycoplasma mycoides

J. Craig Venter Institute
Chapter 1
Origins

It would be easy for us to assume that bacteria are the simplest

form of life and thus presumably would have been the original life
form on this planet. This may be true but it is not a simple equation.
Bacteria are single-cell organisms and are what are known as
prokaryotic cells. These differ considerably from the cells of both
animals and plants inasmuch as there are no visible discrete
compartments within the cell. They are also usually considerably
smaller than the cells of animals and plants.

So how did bacteria first emerge? This looks like a classic ‘chicken
and egg’ conundrum. Bacteria, like all cells, contain DNA and they
function by the decoding of this DNA into proteins, which comprise
the enzymes that control all the major processes within the
organism. In this respect, they are similar to other cells and thus
probably have a common origin. The link between DNA decoding
and protein production is RNA. RNA does not differ greatly in
structure from DNA and some believe that RNA is the origin of life.
This is plausible as RNA is the messenger; it is the molecule that is
transcribed from DNA and from which protein is translated. It was
the discovery of ribozymes by Thomas Cech, at the University of
Colorado, and Sydney Altman of Yale University that strongly
suggested that RNA was the origin of life. Ribozymes are RNA
molecules that have a 3D (tertiary) structure and they can act as
catalysts, similar to enzymes. Therefore RNA could act not only as
the store of genetic material but also as the ‘enzyme’ that decodes it
into the structures of life.

We may never be able to confirm this hypothesis but if we assume

that it is plausible, then we can start to examine how bacteria
emerged and where they fit in the evolutionary tree. Approximately
4.3 billion years ago, the first cells are thought to have arisen,
probably with RNA as an essential catalytic role and later as a self-
replicating molecule. The basic integrity of a cell is the formation of
a cell membrane, composed of lipid bilayers. As these can form
spontaneously, they could have surrounded early RNA molecules.
Their continued presence may have been promoted through
mutation of the RNA, which would have been passed on to
succeeding generations through self-replication. This basic system
does have significant disadvantages because a mistake made in the
replication of RNA would immediately have an effect not only on the
replication of the genetic material but also on the ability to act as a
catalyst—largely, it may be assumed, in a detrimental manner. The
separation of the self-replicating machinery from the enzymes they
encode would have resulted in far fewer abortive stages.
Consequently, we must assume that DNA largely took over the role
of the carrier for the self-replicating genes and proteins of the
enzymes that they eventually encoded. RNA merely remained as the
messenger that carried the instructions from DNA to the formation
of the proteins.

The early bacteria emerged approximately 1.5 billion years after the
creation of the planet. For the next three billion years, bacteria were
probably the planet’s sole living inhabitants. The fossil record shows
that there were huge numbers of bacteria often collecting in large
colonies, attached to many surfaces; their imprint can still be seen.
Approximately one billion years ago, however, the numbers of these
colonies in the fossil record began to fall and this could be evidence
that bacteria had become the source of food for some other life
form. Certainly, this was before the Cambrian explosion of half a
billion years ago, when the diversity of life forms increased rapidly,
but it does suggest that, for the first three-quarters of life on Earth,
bacteria had it all their own way.

What were these bacteria?

The common view is that prokaryotic cells, such as bacteria, and
eukaryotic cells, such as those that comprise our bodies, had a
common ancestor. The last universal common ancestor (LUCA), or
cenancestor, is considered to be the most recent ancestor of all life
on Earth and possibly was living some 3.5 billion years ago (Figure
1). It is thought to have been a prokaryotic, single-celled organism
possibly similar to simple bacteria found today. By this time, it has
been concluded that the genetic code must have become DNA rather
than RNA, and that the catalysts had become true enzymes
(proteins) composed of the 20 amino acids. Furthermore, the
machinery for dividing the DNA, maintaining its integrity, and
expressing the genes through RNA was already established.
1. Timescale of bacteria emergence.

These bacteria would have been exclusively anaerobic; they did not
respire oxygen as there was little or no available oxygen in the
atmosphere at the time. These bacteria could produce energy from
the available nutrients but this was an extremely inefficient process.
About 3.2 billion years ago, photosynthetic bacteria or cyanobacteria
first emerged. These bacteria could use energy from the sun to
make sugars, which were used for further metabolism. The by-
product of this photosynthesis was oxygen, which began to
accumulate in the atmosphere. Oxygen is toxic to many cells,
particularly the early anaerobic bacteria, which probably started to
decline. About 2.5 billion years ago, the fossil record shows that
aerobic bacteria emerged, able to use the newly available oxygen to
convert sugars into energy, usually in the form of adenosine-5´-
triphosphate (ATP). The use of oxygen vastly increased the energy
obtained from a single sugar molecule, and these bacteria soon
predominated.
The main eukaryotic cells may have derived from an early example
of Archaea, which themselves derived from LUCA. They probably
evolved about two billion years ago. These were distinguishable from
prokaryotic cells by having a defined nucleus, usually comprising
discrete chromosomes, that contained the DNA. However, having
emerged from prokaryotes, the evolution of these eukaryotic cells
did not proceed independently of prokaryotes, but with a degree of
symbiosis. Eukaryotic cells possess distinct organelles; most animal
cells possess mitochondria and most plant cells contain chloroplasts.
Both these organelles are approximately the same size as bacteria
and possess their own DNA. Mitochondria probably originated from
oxygen-utilizing bacteria, such as early examples of proteobacteria
or cyanobacteria, which have been captured by eukaryotic cells to
provide energy through oxidative phosphorylation. Chloroplasts
probably originated from photosynthetic bacteria in order to produce
energy from light. In both cases, the eukaryotic cells ingested the
bacteria but did not destroy them; allowing coexistence known as
endosymbiosis.

Both animal and plant eukaryotic cells were taking up the energy-
generating machinery of bacteria, which had evolved over millions of
years, thus obviating the need for their separate evolution in
eukaryotic cells. This certainly accelerated the development of
eukaryotic cells. It is believed that the incorporation of mitochondria
and chloroplasts into eukaryotic cells also occurred 1.5 billion years
ago as eukaryotic cells emerged. The mitochondria-containing cells
became the cells of animals and the chloroplast-containing cells
those of plants. Both mitochondria and chloroplasts still contain their
ancient DNA, which replicates independently from the nuclear DNA
of the eukaryotic cell itself. Similarly, this DNA is not affected by the
sexual reproduction of the host and carries unaltered DNA from
generation to generation. The acquisition of these energy-producing
organelles rapidly increased the evolution of the eukaryotic cells,
resulting in the Cambrian explosion of multicellular, eukaryotic
animals and plants 500 million years ago.
What else distinguishes a bacterial cell from a eukaryotic cell? The
size is an obvious distinction: the bacterial cell could be two microns
in length whereas a mammalian cell may be 15 times longer and
1,000 times greater in volume. However, one striking feature is the
maintenance of the self-replicating genetic material, DNA. In
bacteria, it is maintained as a single genome, with the genes closely
packed and genes controlling related functions often clustered
together. On the other hand, most eukaryotic cells have their DNA
divided up into several chromosomes and are diploid, receiving one
set of chromosomes from each parent. Therefore the chromosomes
are homologous pairs, basically carrying the same genes in the same
order.

This arrangement can compensate for errors in the DNA as

recombination of a damaged gene with the duplicate intact gene can
rectify the error. The single copy of the bacterial genome means that
if a mistake is made, it is likely to be permanent and often is lethal.
The reason for this difference may be that many eukaryotic cells
form part of a much larger organism where disastrous mutations
could cause major damage, such as cancer. So the duplication may
have evolved to minimize this. Bacteria, on the other hand, although
they are single-celled organisms, are usually not in isolation but form
colonies. Each cell is capable of regenerating the colony. So if, during
DNA division, mutations occur which are lethal for the host cell,
there will be enough members of the colony in which mutations have
not occurred to keep the colony thriving.

From a technical point of view, it has been convenient to categorize

bacteria according to their response to the Gram stain. This
technique developed by Hans Christian Gram in 1884 distinguishes
bacteria with a single membrane (Gram-positive) from those that
have a double membrane and much thicker cell wall (Gram-
negative). The simpler cell membrane structure of Gram-positive
bacteria would suggest that they are the ancestral bacteria;
however, there is some evidence that the opposite may be true.
Whichever is the case, it has proved crucial in the examination of
bacteria to separate them into these two broad classes.
Chapter 2
Evolution

Bacteria and their own fight for survival

Normally bacteria divide by binary fission, whereby a single cell
expands by creating macromolecules that make up the components
of the cell and the cell wall. When the cell reaches approximately
double its size, a septum is formed between the two halves of this
enlarged cell. The septum is essentially part of the cell wall that,
when complete, forms two complete daughter cells, which are able
to separate from each other. In ideal conditions, this cell division can
take place once every 20 minutes; so, in theory, a single bacterial
cell could produce more than 16 million progeny in eight hours.
Although cell division time is short, the time taken to replicate the
DNA of the cell is longer, often twice as long. In order to keep up
with cell division, new replication cycles of DNA are started before
the previous round has finished. Indeed, multiple copies of DNA are
being replicated within a fast-growing culture. Rapid replication of
DNA promotes mutations. Many of these will be lethal but some will
be beneficial. In particular, the mutations have allowed individual
bacterial species increased capability to compete for nutrients, often
to the disadvantage of neighbouring bacteria, resulting in some local
‘warfare’, particularly in environments such as the soil. This led to
the development of ‘weapons’, chemicals known as secondary
metabolites that are excreted by the bacteria. Bacteriocins are
released by bacteria to destroy surrounding bacteria of similar
species, but they are quite limited in their effect. This capability has
been refined by, for example, the Actinomyces, which are able to
release chemicals that kill most bacteria; it is from these that we get
most of our antibiotics.

By many small advantageous mutations, bacteria have been able to

adapt to almost every environment and this has been the primary
method of survival and evolution; however, the evolution of bacteria
has not been without its own struggles. A hundred years ago,
Frederick Twort and Félix d’Hérelle independently discovered small
elements capable of killing bacteria. These were bacterial viruses or
bacteriophages, simply called phages for short (Figure 2). Their
existence had been implied since the earliest medical documentation
which reported that some river waters could cure some bacterial
infections.
2. Structure of a T4 bacteriophage.

The conventional lytic phage, which is composed of DNA surrounded

by a protein coat (Figure 2), follows a cycle; it starts when the
phage attaches to the bacterial cell and injects its DNA. This takes
over the normal replication machinery of the bacteria and results in
the production of many, sometimes hundreds, of new phage protein
particles. These particles pick up the replicated phage DNA; they are
released, killing the host bacterial cell in the process, and are able to
infect new bacterial cells. Bacteria have had to defy this attack and,
quite quickly in some cases, the bacteria become resistant, often by
alterations in the cell surface so that the phage cannot attach. There
has been a continuous struggle between bacteria and these viruses
but, as with viruses that attack humans, a balance is usually
reached.

At some points in this process, there would have been an inevitable

exchange of genes. The new phage particles may pick up some
bacterial genes along with, or rather than, new phage DNA which
they can pass on to other bacterial cells on subsequent infection.
Some phages can become parasitic after infection; instead of
producing new phages, the DNA integrates into the bacterial DNA. In
this case, the phage DNA is replicated every time the bacterial DNA
is replicated and so every daughter-cell DNA carries a copy of the
phage DNA. It is often kept integrated by a repressor protein
encoded by the DNA itself. This is known as a lysogenic phage. If
the repressor protein is compromised by a chemical, ultraviolet light,
or some other insult, then the phage DNA initiates the production of
new phage particles, the DNA replicates and is incorporated into
these particles, and they are released to invade other bacteria.

Bacteria and their place in the world

Bacteria are, of course, still evolving; however, if we move out of
geological time and into a time frame with which we are more
familiar, the role of bacteria appears more static. They currently have
an important role to play within some of the biogeochemical cycles
that allow other living organisms to survive; these include the
carbon, nitrogen, and sulphate cycles.

Carbon is the essential element in all living matter and it has to be

converted into different forms. Carbon dioxide, produced by animals
and as a by-product of industry, would soon render the world
uninhabitable; plants fix carbon dioxide but so do bacteria. These
bacteria, known as autotrophs, can produce their own organic
compounds using the sun’s radiation and carbon dioxide and water.
They convert them to sugars by photosynthesis. These can be used
either by the bacteria themselves or by other organisms. Some
carbon is taken up as methane and there are methane-oxidizing
bacteria that convert it to carbon dioxide.

Bacteria are also primarily responsible for the removal of carbon

from dead tissue and plants. On the other hand, some bacteria can
use organic carbon, particularly sugars, for the production of energy
with the release of carbon dioxide. So different bacterial species are
on opposite sides of the cycle and the success of each will depend
on the availability of the nutrients they require. If there is an
increase in carbon dioxide, the photosynthetic bacteria will prosper,
whereas if too much has been converted into sugar, and there is an
abundance of oxygen, respiring bacteria will thrive. The burning of
fossil fuels and the high quantities of carbon dioxide it produces
appear to upset this balance.

Nitrogen is an important component of proteins and nucleic acids,

such as DNA. Most nitrogen on Earth is available as a gaseous
molecule, comprising two nitrogen atoms, in the atmosphere, but
this is relatively inaccessible for plants, and thus eventually
ourselves. The atmospheric nitrogen has to be ‘fixed’ and this is
mainly performed by bacteria that contain a nitrogenase enzyme
that can break the bond between the nitrogen atoms. These atoms
can be converted to an ammonium salt or nitrates, which are usable
by plants. Indeed, the nitrogen-fixing bacteria are often found
clustered around the roots of plants. Bacteria are used for most of
the interconversions of the various forms of nitrogen. Some bacteria,
such as Pseudomonas species, are even able to convert nitrite back
into molecular nitrogen, which is released into the atmosphere, so
an equilibrium is established. This balance is upset when large
quantities of nitrates are used as fertilizers and there are insufficient
bacteria to convert the excess back into atmospheric molecular
nitrogen, resulting in pollution.

Sulphur is an essential component of proteins and some co-factors.

Plants require sulphates but this sulphur derivative is often not
readily available. Sulphur is abundant as inorganic sulphides and
thiosulphates, which can be oxidized by the bacterial genus
Thiobacillus to form sulphates. These can be used directly by plants
and incorporated into proteins that, in turn, can become
incorporated into animal proteins. On the other hand, the balance
can be maintained by anaerobic bacteria, such as the genus
Desulfovibrio, that are able to reduce sulphates to hydrogen
sulphide, which can then be oxidized to elemental sulphur.

Bacteria and the biochemical machinery for all living

cells
Bacteria were the only living cells for more than two billion years and
evolved enormous genetic diversity. During that time, bacteria
developed all the biochemical machinery required for life. They
evolved the use of DNA as the carrier of genetic information, the use
of messenger RNA to convey that information, and the use of
ribosomes and L-amino acids for the production of proteins and thus
the enzymes essential for life. Most of this machinery is identical in
all eukaryotic cells, the complex cells found in plants and animals, so
it is likely that these cells evolved directly from bacteria developing
this machinery for their own needs. The main adaptation of these
plant and animal cells has been to compartmentalize the cell, with
specialized organelles. However, these structures appear to have
come from prokaryotes; the nucleus from the archaeon, the
mitochondria in animals from Proteobacteria species, and the
chloroplasts in plants from photosynthesizing bacteria—all bringing
their own genetic and biochemical machinery to the evolution of
eukaryotic cells.

Bacteria and the evolution of herbivores

Bacteria must have had a major impact on the development of all
species and not just because of their capability to cause disease. As
small eukaryotic animals emerged, they fed on bacteria, indeed
many still do. As animals evolved into multicellular organisms, they
exploited bacteria to their evolutionary gain. Many bacteria have
lived in a symbiotic relationship with other organisms to the
evolutionary advantage of both. Grasses emerged after the
Cretaceous extinction, becoming widespread as the Earth became
drier, to provide probably the most abundant worldwide food
resource, but many animals could not digest it because grass
contains fibrous cellulose. Some animals, such as cattle, evolved a
digestive system that relied upon bacteria. Their stomachs are made
up of four compartments.

The grass enters the rumen, which is essentially a large

fermentation chamber. There are billions of bacteria there, including
the species Ruminococcus flavefacians, Ruminococcus albus,
Bacteroides succinogenes, Butyrivibrio fibrisolvens. These bacteria
break up the fibre of grass cells. Cattle then regurgitate to grind the
products with their teeth (chewing the cud) to try to break up the
fibre further. The bacterial digestion of the grass releases nutrients
that are, of course, used by the bacteria themselves but the excess
is available to the animals as it is passed on to the abomasum, which
contains acid. The nutrients are further broken down and absorbed
by the small intestine. Without bacteria, grass could not be a food
source. The exploitation of bacteria is not restricted to large grass-
eating herbivores. Cockroaches, termites, and many other smaller
animals have been able to expand the variety of plant material on
which they can feed as they acquire new bacteria that are able to
metabolize and release the necessary nutrients.

Bacteria and the evolution of complex organisms

It is now accepted that multicellular organisms and their bacteria do
not evolve separately but maintain an evolutionary relationship. The
holobiont (host plus bacteria) with its hologenome (host genome
plus microbiome) is a level of selection in evolution. as according to
the hologenome theory, the progeny receives genes from its parents’
DNA but bacteria are also passed from parent to progeny. Changes
in the progeny genome may affect the composition of the colonizing
bacteria, which in turn may influence the survival potential of the
host. This bacterial co-evolution with the host has been exploited to
study animal evolution. The microbiota of brown, black, and polar
bears has shown that they diverged far earlier than had been
determined from their own DNA analysis. Bacterial co-evolution may
also have affected the emergence and evolution of Homo sapiens.
An example of recent evolution is the ability of Japanese people to
digest agar, unlike other human populations. The Japanese have a
bacterium in their gut that contains a gene that codes for the
enzyme agarase, which other ethnic groups lack. The agarase gene
was transferred from a marine bacterium, present on raw seaweed,
which forms part of the traditional Japanese diet, to a resident gut
bacterium.
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