Journal of Applied Microbiology 2001, 91, 597±601
The life and works of Louis Pasteur
M. Schwartz
Institut Pasteur, Paris, France
749/1/01: received and accepted 21 February 2001
Being in the UK, I will start by quoting one of the most
famous physicians in the history of this country, the surgeon
Joseph Lister. Addressing Louis Pasteur, he expressed
himself as follows:
It is my great privilege to convey to you, tributes, thanks
and respect from all involved in medicine and surgery; it is
true to say that, of all people in the world today, medical
sciences owe you the most ¼ For centuries, infectious
diseases have been shrouded, as it were under a dark
curtain. In discovering the microbial origin of disease you
have raised that dark curtain! (Lister 1893).
Lister stated this at a ceremony held at the Sorbonne, in
Paris, on the occasion of the 70th birthday anniversary of
Louis Pasteur, on December 27, 1892. On that jubilee
occasion, delegates from throughout the world gathered to
pay homage to the scientist and to express their gratitude
and admiration on behalf of their respective countries.
Joseph Lister represented the Royal Societies of London and
Edinburgh.
In 1892, the thoughts that were uppermost in the minds of
Joseph Lister and the other participants at the jubilee
ceremony no doubt centred on Pasteur's most recent
accomplishments in the ®eld of infectious diseases. Today,
more than a century later, we are far enough removed from
those events to be able to more fully appreciate the impact of
his body of work.
From among Pasteur's early period, in the late 1840s, the
image which comes most readily to mind is that of the young
26-year-old chemist, fresh out of the Ecole Normale, sorting
out crystals of tartaric acid while the eminent crystallogra-
pher, Jean-Baptiste Biot, looked on in astonishment. After
the experiment was over, and according to Pasteur himself,
the illustrious old man was deeply moved. `My dear boy', he
said taking Louis by the arm, `I have loved science so much
all my life that this touches my heart.'
What was this dramatic discovery?
What Pasteur had shown to Biot was that a rare form of
tartrate, referred to as paratartrate, was in fact composed of
equal quantities of two types of molecules whose crystals,
Correspondence to: Maxime Schwartz, Institut Pasteur, 25, rue du Dr. Roux,
75724 Paris Cedex 15 ± France.
ã 2001 The Society for Applied Microbiology
although very similar, could be distinguished by the
orientation of one tiny facet.
In addition to this minute difference in the shape of their
crystals, these two molecular species, though identical in all
of their physical and chemical properties, nonetheless had
one other difference: their solutions rotated the plane of
light polarization in opposite directions. Pasteur hypothes-
ized that these two forms of tartrate corresponded to two
different spatial con®gurations of the atoms within the
molecule, two forms which were asymmetrical in them-
selves, but symmetrical with respect to one another, just as
our two hands are. This was a most revolutionary hypothe-
sis: two molecules, containing the same atoms, linked to one
another by the same bonds, could nevertheless differ in the
spatial arrangement of these atoms. This hypothesis, which
would not be de®nitely con®rmed until 30 years later with
the establishment of the principle of carbon asymmetry, laid
the foundation for stereochemistry. The far-reaching con-
sequences of this concept can only fully be appreciated by
recalling the fact that, as Pasteur himself sensed and as
molecular biology would demonstrate many years later, all
interactions between biological molecules, and hence all life
processes result from the precise three-dimensional arrange-
ment of the atoms within these molecules. Moreover, the
emergence of the critical notions of symmetry and sym-
metry-breaking were of capital importance, for they laid the
basis for a number of modern theories of physics such as that
of elementary particles and phase changes.
Upon observing that all compounds whose solutions
rotated the plane of polarized light (optically active) arose
from plant and animal sources, Pasteur began to suspect that
asymmetry was a sign of life. With this idea in mind, he,
began studies on fermentation in 1854. Fermentation was
known even in ancient times, for it was used in the
preparation of bread, wine and many other types of food and
drink. But when Pasteur began to examine it, the question of
fermentation was the subject of utmost confusion. While in
some cases, the participation of micro-organisms in such
transformations of organic matter vas recognized, in general
their role was completely misunderstood. Justus Liebig, the
renowned German chemist, a contemporary and adversary
of Pasteur, had stated: `Yeast from the malt ¼ (transfers) its
own state of decomposition to that which is around it. The
598 M . S C H W A R T Z
Fig. 1 Louis Pasteur
movement that disturbs the balance imprinted in its own elements
also communicates with other elements of bodies in contact with
it' (Liebig 1839).
Needless to say, Pasteur could not accept such a confused
explanation. The observation that optically active products
appeared during the process of fermentation led him to
propose an entirely different and much more accurate
hypothesis. To Pasteur, `¼ fermentation, far from being a
lifeless phenomenon, is a living process ¼ all phenomena of
fermentation correlate with the development of mycodermic cells
and plants which I have prepared and studied in an isolated and
pure state'. Indeed, he describes this method in his Note on
lactic fermentation, published in 1857 and which can be
considered as the birth certi®cate of Microbiology.
The purity of a ferment, its homogeneity, its unhindered
development, with the help of a nutrient perfectly adapted
to its individual nature, this is one of the essential conditions
for obtaining high-quality fermentations (Pasteur 1857).
These are the basic principles of microbiology: isolate a
micro-organism, and provide it with the adequate growth
medium. With these simple rules, man had learned to
master the microbe. As applied by Pasteur and his students
to the preparation of wine, beer, vinegar and dairy products,
these principles were to revolutionize those industries. They
formed the basis for what we now refer to as biotechnology.
Each fermentation process results from the action of a
speci®c micro-organism. To obtain a good fermentation, one
ã 2001 The Society for Applied Microbiology, Journal of Applied Microbiology, 91, 597±601
has to introduce the right kind of micro-organism, or, at
least, make sure that it is present, and avoid the presence of
other types of micro-organisms, which could alter the
process. Once the product is made, contaminating micro-
organisms could still alter it. It is to prevent this alteration,
in the case of wine, that Pasteur introduced a process now
named after him, pasteurization.
Before Pasteur, when milk was getting sour, when meat or
®sh were decaying, nobody really knew why. Pasteur
demonstrated that putrefaction, like fermentation, was due
to the growth of micro-organisms. The empirical methods
used in the past to delay food decay could then be
rationalized and improved.
Pasteur understood that putrefaction, unpleasant as it may
be, plays a major role in the recycling of elements between
the living world and the mineral world. This `immense role of
in®nitely small bodies in the general economy of nature', as
Pasteur himself says (Pasteur 1862), warrants re¯ection in
our present day era in which the protection of environment
has become one of humanity's major priorities.
Fermentation and putrefaction were often perceived as
being spontaneous phenomena. But what was Pasteur to
make of the ferments and yeasts which he felt were essential
in the unfolding of those processes? Where did they come
from? Did they appear spontaneously in the media, or did
they come from elsewhere? Pasteur could not escape this
debate, a debate stemming from ancient beliefs, concerning
the spontaneous generation of bees, frogs, mice, etc., but
which had taken on new strength toward the end of the 17th
century following the discovery of `animalcules' by Anton
van Leeuwenhoek, the Dutch inventor of the microscope.
On the subject of these microscopic organisms, the 18th
century witnessed such supporters of spontaneous genera-
tion as the English scholar John Needham and France's
renowned naturalist Buffon taking part in heated debate
with other scientists such as the Italian abbot Spallanzani.
Pasteur kept an open mind in his approach to this question.
In 1859, in a letter to the man who was to become his most
ardent opponent on the subject, Felix Archimede Pouchet,
he wrote that the question of spontaneous generation was
`entirely open and still awaiting proof ' and that all of this was
`unknown and warranted experimentation' (Pasteur 1859).
Pasteur responded to his own invitation. By extremely
painstaking experimental methodology, he demonstrated
that the appearance of micro-organisms in a presterilized
medium could always be explained by germs coming from
the outside. He thus succeeded in discrediting any experi-
mental basis for the theory of spontaneous generation.
On a philosophical level, the repercussions were resound-
ing. The onset of life was decidedly not a predictable
phenomenon, regularly occurring in any fermentable
medium. The question of the origin of life was thereafter
clearly set forth ± and remains so today.

Pasteur's research on fermentation and on so-called
spontaneous generation inevitably led him toward the study
of infectious diseases. He had only recently succeeded in
demonstrating that if environmental yeasts are prevented
from being deposited on grapes, the juices of these grapes
will not ferment, when he wrote:
`By analogy, is it unreasonable to hope that the day will
come when easily administered preventive measures will be
able to stop the scourges which terrify and decimate
populations, such as yellow fever and the bubonic plague?'
(Pasteur 1879).
In other words, infectious diseases, like fermentations, are
probably due to `germs', and it may be possible to protect
human beings against them, as one can `protect' grapes
against yeast.
But it was neither yellow fever nor bubonic plague that
Pasteur would attempt to cure; that task would be left up
to his followers. Instead, upon the request of his former
mentor who was now senator in the department of the
Gard, Jean-Baptiste Dumas, he next turned his attention
toward silkworm disease. Pasteur knew virtually nothing
about silkworms, but he accepted the challenge, seizing the
opportunity to learn more about infectious diseases. After
5 years of painstaking research, from 1865 to 1870, he
succeeded in saving the silk industry through a method
called `graining' which enabled the preservation of healthy
eggs while eliminating those from contaminated females.
This method is still used today in silk-producing countries.
Indeed, in Japan, a law actually exists which controls its
use!
As we mentioned earlier, according to Joseph Lister, it
was Pasteur who demonstrated the microbial nature of
infectious diseases. However, this does not imply that
Pasteur alone was responsible for identifying the microbes
which caused all major diseases. He did, of course,
contribute to de®nitively identifying the agent of anthrax,
which was decimating cattle and sheep herds; he identi®ed
staphylococcus, streptococcus and the septic vibrion. But
Robert Koch, of the German school, provided an equally
important contribution.
In reality, what is so uniquely important about the entire
body of Pasteur's work is that it laid the groundwork upon
which the microbial theory of disease was built.
He demonstrated how to cultivate bacteria and later, for
rabies, he set forth the premizes for the culture of viruses on
animal tissues.
He had always been intrigued by the mechanisms through
which pathogenic microbes caused profound disturbances in
the physiology of the infected organism, what we would call
today the mechanisms of pathogenicity. Not surprisingly, it
is one of his closest collaborators, Emile Roux, who made
the ®rst major contribution to this ®eld, by identifying
diphtheria toxin.
ã 2001 The Society for Applied Microbiology, Journal of Applied Microbiology, 91, 597±601
THE LIFE AND WORKS OF LOUIS PASTEUR 599
Pasteur also demonstrated how pathogens are able to
spread through animal and human populations, thereby
laying the foundation for infectious epidemiology and
de®ning the basic rules of hygiene.
Concerning epidemiology, I would like to recall two facts,
one well known and one which is less, concerning anthrax.
The well known story is that of the so-called `cursed
®elds', where it seemed that the cattle would systematically
catch anthrax, and die. One day, while Pasteur was having a
walk close to one such ®eld, he noticed that the ground had a
different colour in one place. He was told that, the year
before, sheep which had died of anthrax had been buried at
this spot. Getting closer, Pasteur noticed the presence, at the
surface, of a multitude of those twisted tubes of earth which
are excreted by worms. The idea then came to him that the
worms, travelling up and down in the ground, could bring to
the surface anthrax spores present in the dead bodies. He
veri®ed that, indeed, the excrements of worms taken at this
place contained spores of the anthrax bacillus. The cattle
could then contaminate itself by grazing in such spots. The
cursed ®elds were thereafter eliminated, once farmers ceased
to bury the dead animals in them.
Less well known, perhaps, are observations made, in 1881
by Ed. Nocard a veterinarian and a collaborator of Pasteur.
At that time the battle was still raging between the
`contagionists' and the `spontaneists'. The former, with
Pasteur, contended that infectious diseases could only be
caught through contagion, whereas the latter claimed that
they could appear spontaneously. In that context, Nocard
was informed of the appearance of anthrax in farms where it
had never been seen before, and where it thus must have
appeared `spontaneously'. I will only relate his observations
concerning one of these farms.
There, a young farmer full of dynamism, had decided to
improve the yield of his ®elds. With this in mind he bought
a large quantity of fertilizer, something unheard of in that
area. He then got beautiful crops but, the year after, when he
let his cattle graze on the same ®elds a large number
immediately caught anthrax. That year he lost about a
quarter of his herd. Nocard then discovered that the
fertilizer was commercialized by a company that collected
the carcasses of dead animals over a large area and did not
take special precautions in transforming them into fertil-
izers. Among the carcasses some were bound to have been
from animals which had died of anthrax, and the spores they
contained had thus been distributed in the ®eld of the
unlucky young farmer. I am sure you will have already
drawn a parallel with what happened at a time much closer
to us in the dissemination of BSE.
Going back to Louis Pasteur himself, the last part of his
life was no less productive than the rest. It is then, indeed,
that he outlined the overall principles of vaccination and
contributed to the foundation of immunology.
600 M . S C H W A R T Z
The concept of acquired immunity dates from ancient
times. Indeed, Thucydides reported that those who were
cured of the plague no longer ran the risk of falling victim to
the disease. The ®rst vaccine was developed by the English
physician, Edward Jenner, who, at the end of the 18th
century discovered that human beings could be protected
against smallpox by inoculation of the similar but benign
disease, called `cowpox' in English, and `vaccine' in French.
Although of crucial importance, since it lead to complete
eradication of smallpox in 1979, this empirical discovery
could not be generalized to other diseases.
It is often said that Jenner discovered vaccination, and
Pasteur invented vaccines.
Pasteur's fundamental discovery in this ®eld dates back to
1879 and concerned the disease known as fowl cholera,
which was rampant in chicken coops at that time.
The disease was due to bacteria which today bear the
name `Pasteurella'. When inoculated into a chicken, several
drops of a culture of these bacteria were suf®cient to kill the
animal. But Pasteur noted, partly by chance it would seem,
that chicken inoculated with an old culture not only did not
die but were protected against a later inoculation with a
virulent culture. The principle of vaccination with attenu-
ated pathogens was thus discovered.
From then on, Pasteur repeatedly applied this principle to
other diseases. His ®rst great success both at a scienti®c
1evel and in terms of public opinion ± today we would refer
to it as a `mass media happening' ± was the vaccination
against anthrax. The famous public experiment held in
Pouilly le-Fort in 1881, during the course of which
24 vaccinated sheep survived an injection of the anthrax
bacillus, while 24 nonvaccinated sheep died, had extra-
ordinary repercussions, convincing a large portion of public
opinion of the validity of Pasteur's work.
The few remaining sceptics rallied around Pasteur for his
®nal victory, that of human vaccination against rabies. The
problem was complicated from the very outset, for the rabies
microbe was invisible ± we now know that it is a virus rather
than a bacterium ± and could not multiply in any culture
medium. But the stakes were high, for although the disease
was relatively rare in France, it had always fascinated the
popular imagination, conjuring up fear and mystery. For
Pasteur, conquering rabies would consolidate the ®nal
victory of his theories.
Even though he could not see or cultivate the microbe,
Pasteur knew that it had to be there. It had to be in the
nervous system, recognized as its target. As a replacement
for in vitro cultivation, Pasteur transmitted the infectious
agent from animal to animal, by intracerebral inoculation.
He adapted the disease to the rabbit, and then undertook to
attenuate the invisible microbe, which he did by dessication
of the spinal cord of infected animals. In doing so, he
thought, at ®rst, that he was doing the same as with chicken
ã 2001 The Society for Applied Microbiology, Journal of Applied Microbiology, 91, 597±601
cholera and anthrax, i.e. create an attenuated form of the
microbe. In fact, as he was to realize later, most of the virus
was presumably killed in his preparations. Thus, rather
unknowingly, he opened the way for the second class of
vaccines, besides attenuated live micro-organisms, and
consisting in inactivated micro-organisms, which would
themselves later lead to subunit vaccines.
The story of the vaccination against rabies is well known.
So well known, in fact, that for many this was the sole
accomplishment of Pasteur!
The success of the vaccination of Joseph Meister on July 6,
1885 and of the shepherd boy Jean-Baptiste Jupille in
October of that same year, followed by hundreds of other
bite-victims from throughout the world, brought glory to
Pasteur and opened up the era of preventive medicine. On
the heels of this success, the `Academie des Sciences'
launched an international fund-raising campaign to build the
Institut Pasteur. It was there that Louis Pasteur lived out his
last years, it is there that he lies in his ®nal resting place, and
it is there that, for over a century now, the work that he
began is being pursued by his followers.
Louis Pasteur has left us many messages, which I have
tried to brie¯y summarize in this presentation. In ending,
I would like to quote him once more, and leave you with a
last message, which seemed to me of particular relevance at a
time when we are confronted with such terrifying emerging
infectious diseases, as are AIDS and BSE. This quote is
from an article published in 1881, the year of the vaccine
against anthrax, when Pasteur had just found that virulence
is not a ®xed trait of micro-organisms. Virulence could be
weakened to create attenuated strains, why could it not be
increased under other conditions? Here is the quotation:
And so it is that virulence appears to us in a new light,
rather disquieting for humanity, unless nature in its
evolution during centuries of the past already encountered
all possible occasions of creating virulent or contagious
diseases, something which is very unlikely.
What makes a micro-organism harmless for a human being or
any given animal? It is a micro-organism which cannot grow
in our body or in the body of this animal; but nothing proves
that provided this micro-organism were to penetrate one of
the thousands of species of Creation, it might not invade it
and make it ill. Its virulence, then reinforced by successive
passages through members of this species, could become able to
infect some animal of large size, man or certain domestic
animals. In this way new virulences or contagions could be
created. (Pasteur 1881)
REFERENCES
Liebig, J. (1839) Sur les pheÂnomenes de la fermentation et de la
putreÂfaction, et sur les causes qui les provoquent. Annales de Chimie
et de Physique. 2e Serie LXXI, 178.

Lister, J. (1893) In Jubile de Louis Pasteur, Paris, 1893. pp. 16±17. Paris:
Gauthier-Villars et Fils.
Pasteur, L. (1857) Memoire sur la fermentation appeleÂe lactique.
In Memoires de 1a SocieÂteÂdes Science, de l ¢Agriculture et des arts de
Lille, seÂance du 3 aouÂt 1857, 2e SeÂrie, V, pp. 13±26.
Pasteur, L. (1859) Lettre de Pasteur aÁ Pouchet, Paris 28 feÂvrier.
Archives du Museum d'Histoires Naturelles de Rouen, no. 1023 du
catalogue de la Bibliotheque.
ã 2001 The Society for Applied Microbiology, Journal of Applied Microbiology, 91, 597±601
THE LIFE AND WORKS OF LOUIS PASTEUR 601
Pasteur, L. (1862) Note remise au MinisteÁre de l'Instruction publique et des
cultes, sur sa demande, Avril 1862. In L'oeuvre de Pasteur, T.VII, p. 3.
Pasteur, L. (1879) Examen critique d'un eÂcrit posthume de Claude Bernard
sur la fermentation. Paris: Gauthier-Villars.
Pasteur, L. (1881) De l¢atteÂnuation des virus et de leur retour a la
virulence (avec la collaboration de MM.Chamberland et Roux).
Comptes rendus de l¢AcadeÂmie des Sciences, SeÂance du 28 Fevrier XCII,
429±435.