Germ theory, in medicine, the theory that certain diseases are caused by the invasion of the body by

microorganisms, organisms too small to be seen except through a microscope. The French chemist and
microbiologist Louis Pasteur, the English surgeon Joseph Lister, and the German physician Robert Koch
are given much of the credit for development and acceptance of the theory. In the mid-19th century
Pasteur showed that fermentation and putrefaction are caused by organisms in the air; in the 1860s
Lister revolutionized surgical practice by utilizing carbolic acid (phenol) to exclude atmospheric germs
and thus prevent putrefaction in compound fractures of bones; and in the 1880s Koch identified the
organisms that cause tuberculosis and cholera.

Louis Pasteur (born December 27, 1822, Dole, France—died September 28, 1895, Saint-Cloud) French
chemist and microbiologist who was one of the most important founders of medical microbiology.
Pasteur’s contributions to science, technology, and medicine are nearly without precedent. He
pioneered the study of molecular asymmetry; discovered that microorganisms cause fermentation and
disease; originated the process of pasteurization; saved the beer, wine, and silk industries in France; and
developed vaccines against anthrax and rabies.

Pasteur’s academic positions were numerous, and his scientific accomplishments earned
him France’s highest decoration, the Legion of Honour, as well as election to
the Académie des Sciences and many other distinctions. Today there are some 30
institutes and an impressive number of hospitals, schools, buildings, and streets that
bear his name—a set of honours bestowed on few scientists.
Early education
Pasteur’s father, Jean-Joseph Pasteur, was a tanner and a sergeant major decorated
with the Legion of Honour during the Napoleonic Wars. This fact probably instilled in
the younger Pasteur the strong patriotism that later was a defining element of his
character. Louis Pasteur was an average student in his early years, but he was gifted in
drawing and painting. His pastels and portraits of his parents and friends, made when
he was 15, were later kept in the museum of the Pasteur Institute in Paris. After
attending primary school in Arbois, where his family had moved, and secondary
school in nearby Besançon, he earned his bachelor of arts degree (1840) and bachelor of
science degree (1842) at the Royal College of Besançon.

Research career of Louis Pasteur

Louis Pasteur and fermentation
French chemist and microbiologist Louis Pasteur made many important contributions to science, including the
discovery that microorganisms cause fermentation and disease.
In 1843 Pasteur was admitted to the École Normale Supérieure (a teachers’ college in
Paris), where he attended lectures by French chemist Jean-Baptiste-André Dumas and
became Dumas’s teaching assistant. Pasteur obtained his master of science degree in
1845 and then acquired an advanced degree in physical sciences. He later earned his
doctorate in sciences in 1847. Pasteur was appointed professor of physics at the Dijon
Lycée (secondary school) in 1848 but shortly thereafter accepted a position as professor
of chemistry at the University of Strasbourg. On May 29, 1849, he married Marie
Laurent, the daughter of the rector of the university. The couple had five children;
however, only two survived childhood.
Molecular asymmetry
Soon after graduating from the École Normale Supérieure, Pasteur became puzzled by
the discovery of the German chemist Eilhardt Mitscherlich, who had shown that
tartrates and paratartrates behaved differently toward polarized light: tartrates rotated
the plane of polarized light, whereas paratartrates did not. This was unusual because
the compounds displayed identical chemical properties. Pasteur noted that the tartrate
crystals exhibited asymmetric forms that corresponded to their optical asymmetry. He
made the surprising observation that crystalline paratartrate consisted of a mixture of
crystals in a right-handed configuration. However, when these crystals were separated
manually, he found that they exhibited right and left asymmetry. In other words, a
balanced mixture of both right and left crystals was optically inactive. Thus, Pasteur
discovered the existence of molecular asymmetry, the foundation of stereochemistry, as
it was revealed by optical activity. Over the course of the next 10 years, Pasteur further
investigated the ability of organic substances to rotate the plane of polarized light. He
also studied the relationship that existed between crystal structure and molecular
configuration. His studies convinced him that asymmetry was one of the fundamental
characteristics of living matter.
Germ theory of fermentation
In 1854 Pasteur was appointed professor of chemistry and dean of the science faculty at
the University of Lille. While working at Lille, he was asked to help solve problems
related to alcohol production at a local distillery, and thus he began a series of studies on
alcoholic fermentation. His work on these problems led to his involvement in tackling a
variety of other practical and economic problems involving fermentation. His efforts
proved successful in unraveling most of these problems, and new
theoretical implications emerged from his work. Pasteur investigated a broad range of
aspects of fermentation, including the production of compounds such as lactic acid that
are responsible for the souring of milk. He also studied butyric acid fermentation.

In 1857 Pasteur left Lille and returned to Paris, having been appointed manager and
director of scientific studies at the École Normale Supérieure. That same year he
presented experimental evidence for the participation of living organisms in all
fermentative processes and showed that a specific organism was associated with each
particular fermentation. This evidence gave rise to the germ theory of fermentation.
Pasteur effect
The realization that specific organisms were involved in fermentation was further
supported by Pasteur’s studies of butyric acid fermentation. These studies led Pasteur to
the unexpected discovery that the fermentation process could be arrested by passing air
(that is, oxygen) through the fermenting fluid, a process known today as the Pasteur
effect. He concluded that this was due to the presence of a life-form that could function
only in the absence of oxygen. This led to his introduction of the
terms aerobic and anaerobic to designate organisms that live in the presence or absence
of oxygen, respectively. He further proposed that the phenomena occurring
during putrefaction were due to specific germs that function under anaerobic
conditions.
Pasteurization
Pasteur readily applied his knowledge of microbes and fermentation to the wine and
beer industries in France, effectively saving the industries from collapse due to problems
associated with production and with contamination that occurred during export. In
1863, at the request of the emperor of France, Napoleon III, Pasteur
studied wine contamination and showed it to be caused by microbes. To prevent
contamination, Pasteur used a simple procedure: he heated the wine to 50–60 °C (120–
140 °F), a process now known universally as pasteurization. Today pasteurization is
seldom used for wines that benefit from aging, since it kills the organisms that
contribute to the aging process, but it is applied to many foods and beverages,
particularly milk.
Find out about beer ingredients and fermentation and the process of brewing beer
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Following Pasteur’s success with wine, he focused his studies on beer. By developing
practical techniques for the control of beer fermentation, he was able to provide a
rational methodology for the brewing industry. He also devised a method for the
manufacturing of beer that prevented deterioration of the product during long periods
of transport on ships.

Spontaneous generation
Fermentation and putrefaction were often perceived as being spontaneous phenomena,
a perception stemming from the ancient belief that life could generate spontaneously.
During the 18th century the debate was pursued by the English naturalist and Roman
Catholic divine John Turberville Needham and the French naturalist Georges-Louis
Leclerc, count de Buffon. While both supported the idea of spontaneous generation,
Italian abbot and physiologist Lazzaro Spallanzani maintained that life could never
spontaneously generate from dead matter. In 1859, the year English naturalist Charles
Darwin published his On the Origin of Species, Pasteur decided to settle this dispute. He
was convinced that his germ theory could not be firmly substantiated as long as belief in
spontaneous generation persisted. Pasteur attacked the problem by using a simple
experimental procedure. He showed that beef broth could be sterilized by boiling it in a
“swan-neck” flask, which has a long bending neck that traps dust particles and other
contaminants before they reach the body of the flask. However, if the broth was boiled
and the neck of the flask was broken off following boiling, the broth, being reexposed to
air, eventually became cloudy, indicating microbial contamination. These experiments
proved that there was no spontaneous generation, since the boiled broth, if never
reexposed to air, remained sterile. This not only settled the philosophical problem of the
origin of life at the time but also placed on solid ground the new science of bacteriology,
which relied on proven techniques of sterilization and aseptic manipulation.
Work with silkworms
In 1862 Pasteur was elected to the Académie des Sciences, and the following year he was
appointed professor of geology, physics, and chemistry at the École des Beaux-
Arts (School of Fine Arts). Shortly after this, Pasteur turned his attention
to France’s silkworm crisis. In the middle of the 19th century, a mysterious disease had
attacked French silkworm nurseries. Silkworm eggs could no longer be produced in
France, and they could not be imported from other countries, since the disease had
spread all over Europe and had invaded the Caucasus region of Eurasia, as well as China
and Japan. By 1865 the silkworm industry was almost completely ruined in France and,
to a lesser extent, in the rest of western Europe. Pasteur knew virtually nothing about
silkworms, but, upon the request of his former mentor Dumas, Pasteur took charge of
the problem, accepting the challenge and seizing the opportunity to learn more
about infectious diseases. He soon became an expert silkworm breeder and identified
the organisms that caused the silkworm disease. After five years of research, he
succeeded in saving the silk industry through a method that enabled the preservation of
healthy silkworm eggs and prevented their contamination by the disease-causing
organisms. Within a couple of years, this method was recognized throughout Europe; it
is still used today in silk-producing countries.

In 1867 Pasteur resigned from his administrative duties at the École Normale
Supérieure and was appointed professor of chemistry at the Sorbonne, a university
in Paris. Although he was partially paralyzed (left hemiplegia) in 1868, he continued his
research. For Pasteur, the study of silkworms constituted an initiation into the problem
of infectious diseases, and it was then that he first became aware of the complexities of
infectious processes. Accustomed as he was to the constancy and accuracy of laboratory
procedures, he was puzzled by the variability of animal life, which he had come to
recognize through his observation that individual silkworms differed in their response to
disease depending on physiological and environmental factors. By investigating these
problems, Pasteur developed certain practices of epidemiology that served him well a
few years later when he dealt with animal and human diseases.

Vaccine development of Louis Pasteur

Louis Pasteur
French chemist and microbiologist Louis Pasteur experimenting on a chloroformed rabbit, coloured wood
engraving, 1885.
In the early 1870s Pasteur had already acquired considerable renown and respect
in France, and in 1873 he was elected as an associate member of the Académie de
Médecine. Nonetheless, the medical establishment was reluctant to accept his germ
theory of disease, primarily because it originated from a chemist. However, during the
next decade, Pasteur developed the overall principle of vaccination and contributed to
the foundation of immunology.

Pasteur’s first important discovery in the study of vaccination came in 1879 and
concerned a disease called chicken cholera. (Today the bacteria that cause the disease
are classified in the genus Pasteurella.) Pasteur said, “Chance only favours the prepared
mind,” and it was chance observation through which he discovered that cultures of
chicken cholera lost their pathogenicity and retained “attenuated” pathogenic
characteristics over the course of many generations. He inoculated chickens with the
attenuated form and demonstrated that the chickens were resistant to the fully virulent
strain. From then on, Pasteur directed all his experimental work toward the problem
of immunization and applied this principle to many other diseases.

Pasteur began investigating anthrax in 1879. At that time an anthrax epidemic in France
and in some other parts of Europe had killed a large number of sheep, and the disease
was attacking humans as well. German physician Robert Koch announced the isolation
of the anthrax bacillus, which Pasteur confirmed. Koch and Pasteur independently
provided definitive experimental evidence that the anthrax bacillus was indeed
responsible for the infection. This firmly established the germ theory of disease, which
then emerged as the fundamental concept underlying medical microbiology.

Pasteur wanted to apply the principle of vaccination to anthrax. He prepared attenuated

cultures of the bacillus after determining the conditions that led to the organism’s loss of
virulence. In the spring of 1881 he obtained financial support, mostly from farmers, to
conduct a large-scale public experiment of anthrax immunization. The experiment took
place in Pouilly-le-Fort, located on the southern outskirts of Paris. Pasteur immunized
70 farm animals, and the experiment was a complete success. The vaccination
procedure involved two inoculations at intervals of 12 days with vaccines of different
potencies. One vaccine, from a low-virulence culture, was given to half the sheep and
was followed by a second vaccine from a more virulent culture than the first. Two weeks
after these initial inoculations, both the vaccinated and control sheep were inoculated
with a virulent strain of anthrax. Within a few days all the control sheep died, whereas
all the vaccinated animals survived. This convinced many people that Pasteur’s work
was indeed valid.

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Louis Pasteur’s Contributions to Science

Following the success of the anthrax vaccination experiment, Pasteur focused on the
microbial origins of disease. His investigations of animals infected by pathogenic
microbes and his studies of the microbial mechanisms that cause harmful physiological
effects in animals made him a pioneer in the field of infectious pathology. It is often said
that English surgeon Edward Jenner discovered vaccination and that Pasteur invented
vaccines. Indeed, almost 90 years after Jenner initiated immunization with his smallpox
vaccine, Pasteur developed another vaccine—the first vaccine against rabies. He had
decided to attack the problem of rabies in 1882, the year of his acceptance into
the Académie Française. Rabies was a dreaded and horrible disease that had fascinated
popular imagination for centuries because of its mysterious origin and the fear it
generated. Conquering it would be Pasteur’s final endeavour.
Louis Pasteur
Louis Pasteur, coloured lithograph from Vanity Fair (1887).
Pasteur suspected that the agent that caused rabies was a microbe (the agent was later
discovered to be a virus, a nonliving entity). It was too small to be seen under
Pasteur’s microscope, and so experimentation with the disease demanded the
development of entirely new methodologies. Pasteur chose to conduct his experiments
using rabbits and transmitted the infectious agent from animal to animal by
intracerebral inoculations until he obtained a stable preparation. In order
to attenuate the invisible agent, he desiccated the spinal cords of infected animals until
the preparation became almost nonvirulent. He realized later that, instead of creating an
attenuated form of the agent, his treatment had actually neutralized it. (Pasteur
perceived the neutralizing effect as a killing effect on the agent, since he suspected that
the agent was a living organism.) Thus, rather unknowingly, he had produced, instead of
attenuated live microorganisms, a neutralized agent and opened the way for the
development of a second class of vaccines, known as inactivated vaccines.

On July 6, 1885, Pasteur vaccinated Joseph Meister, a nine-year-old boy who had been
bitten by a rabid dog. The vaccine was so successful that it brought immediate glory and
fame to Pasteur. Hundreds of other bite victims throughout the world were
subsequently saved by Pasteur’s vaccine, and the era of preventive medicine had begun.
An international fund-raising campaign was launched to build the Pasteur Institute in
Paris, the inauguration of which took place on November 14, 1888.
Implications of Pasteur’s work

Louis Pasteur
French chemist and microbiologist Louis Pasteur (1852).
The theoretical implications and practical importance of Pasteur’s work were immense.
Pasteur once said, “There are no such things as pure and applied science; there are
only science and the application of science.” Thus, once he established the theoretical
basis of a given process, he investigated ways to further develop industrial applications.
(As a result, he deposited a number of patents.)

However, Pasteur did not have enough time to explore all the practical aspects of his
numerous theories. One of the most important theoretical implications of his later
research, which emerged from his attenuation procedure for vaccines, is the concept
that virulence is not a constant attribute but a variable property—a property that can be
lost and later recovered. Virulence could be decreased, but Pasteur suspected that it
could be increased as well. He believed that increased virulence was what gave rise
to epidemics. In Louis Pasteur, Free Lance of Science (1950), American
microbiologist René Dubos quoted Pasteur:

Thus, virulence appears in a new light which may be disturbing for the future of humanity
unless nature, in its long evolution, has already had the occasions to produce all
possible contagious diseases—a very unlikely assumption.
What is a microorganism that is innocuous to man or to a given animal species? It is a living
being which does not possess the capacity to multiply in our body or in the body of the animal.
But nothing proves that if the same microorganism should chance to come into contact with
some other of the thousands of animal species in the Creation, it might invade it and render it
sick. Its virulence might increase by repeated passages through that species, and might
eventually affect man or domesticated animals. Thus might be brought about a new virulence
and new contagions. I am much inclined to believe that such mechanisms would explain
how smallpox, syphilis, plague, yellow fever, etc. have come about in the course of time, and
how certain great epidemics appear once in a while.
Pasteur was the first to recognize variability in virulence. Today this concept remains
relevant to the study of infectious disease, especially with regard to understanding the
emergence of diseases such as bovine spongiform encephalopathy (BSE), severe acute
respiratory syndrome (SARS), and acquired immunodeficiency syndrome (AIDS).

Louis Pasteur
A tribute from Le Petit Journal, Paris, at the time of Louis Pasteur's death (1895).
After Pasteur’s 70th birthday, which was acknowledged by a large but solemn
celebration at the Sorbonne that was attended by several prominent scientists, including
British surgeon Joseph Lister, Pasteur’s health continued to deteriorate. His paralysis
worsened, and he died on September 28, 1895. He was buried in the cathedral of Notre-
Dame de Paris, but his remains were transferred to a Neo-Byzantine crypt at the Pasteur
Institute in 1896.
University of Strasbourg
Le Bel Institute, University of Strasbourg, France. From 1971 to 2009, this portion of the school was named
Louis Pasteur University (Strasbourg I).
During Pasteur’s career, he touched on many problems, but a simple description of his
achievements does not do justice to the intensity and fullness of his life. He never
accepted defeat, and he always tried to convince skeptics, though his impatience and
intolerance were notorious when he believed that truth was on his side. Throughout his
life he was an immensely effective observer and readily integrated relevant observations
into conceptual schemes.
Agnes Ullmann
Émile Roux
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Émile Roux
French bacteriologist
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Also known as: Pierre-Paul-Émile Roux
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Émile Roux
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In full:

Pierre-Paul-Émile Roux
Born:

Dec. 17, 1853, Confolens, Charente, France

Died:

Nov. 3, 1933, Paris (aged 79)

Awards And Honors:

Copley Medal (1917)

Subjects Of Study:

diphtheria

tetanus
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Émile Roux (born Dec. 17, 1853, Confolens, Charente, France—died Nov. 3, 1933,
Paris) French bacteriologist noted for his work on diphtheria and tetanus and for his
collaboration with Louis Pasteur in the development of vaccines.

Roux began his medical studies at the University of Clermont-Ferrand. In 1878 he was
accepted into Pasteur’s laboratory at the University of Paris and spent 10 years there,
completing his medical degree in 1881. During that period his work was integral to the
development of methods used in preparing vaccines for diseases such as
fowl cholera, anthrax, and rabies. In 1888 Roux joined the newly created Pasteur
Institute, where, with Alexandre Yersin, he demonstrated that the symptoms of
diphtheria are caused by a toxin secreted by the diphtheria bacillus (a bacterium). That
discovery, together with the subsequent finding by bacteriologists Emil von
Behring and Kitasato Shibasaburo that infection with the diphtheria bacillus elicits the
production of an antitoxin (antibody), led to the development of
diphtheria immunization and serum therapy. Roux became director of the Pasteur
Institute in 1904, a post he held until his death in 1933.
University of Strasbourg
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University of Strasbourg
university, Strasbourg, France
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Also known as: Université de Strasbourg, Universities of Strasbourg I, II, and III
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University of Strasbourg
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French:

Université de Strasbourg
Formerly (1971–2008):

Universities of Strasbourg I, II, and III

Areas Of Involvement:

Protestantism
Notable Alumni:

Jules Hoffmann

Alberto Fujimori

Emmanuel Lévinas

Albert Schweitzer

Jean-Claude Juncker
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University of Strasbourg, autonomous state-financed institution of higher learning
in Strasbourg, France. The original university was founded by Protestants in 1537 as a
German gymnasium (secondary school for the study of the classics) when Strasbourg
was under German rule. The gymnasium became an academy in 1566 and a university in
1621. Under France’s 1968 Orientation Act, which reformed higher education, the
university was separated into three institutions—the Universities of Strasbourg I, II, and
II—in 1971. They were reunited as the University of Strasbourg in January 2009.

The school was influenced from its beginning by the Protestant Reformation and
subsequently by the continuing struggle between France and Germany for control of the
province of Alsace. Along with Strasbourg itself, the university passed into French hands
in 1681. It was suppressed in 1792 during the French Revolution and was succeeded in
the early 19th century by various schools and faculties that were loosely consolidated
into a new university. After the Franco-German War ended in 1871, France ceded
Strasbourg to Germany and the university was reorganized, becoming the Emperor
William University of Strasbourg (1872–1918). In 1919 the city again came under French
control and the university was reestablished as an autonomous French university. From
1939 to 1945, during the German occupation of World War II, the school’s faculties were
moved to Clermont Ferrand University in central France, then back to Strasbourg in
1945.

Johannes Sturm (1507–89), the principal founder, directed the Strasbourg Gymnasium
for more than 40 years. His method of graded courses of studies shaped secondary
school programs throughout Europe. Johann Wolfgang von Goethe completed law
studies at Strasbourg in 1771, and Louis Pasteur was a professor of chemistry
there. Albert Schweitzer studied medicine there from 1905 to 1913.

The modern university is composed of a number of teaching and research units. These
schools, institutes, and faculties specialize in many areas of study, including medicine,
sciences and technology, languages and letters, law, economics, management, political
science, and other social sciences.
This article was most recently revised and updated by Heather Campbell.
fermentation
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fermentation
chemical reaction
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plant producing corn ethanol
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Key People:

Louis Pasteur

Hans von Euler-Chelpin

Sir Arthur Harden

Eduard Buchner
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top fermentation

industrial fermentation

tank fermentation

cap

malolactic fermentation
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Fermentation, chemical process by which molecules such as glucose are broken down
anaerobically. More broadly, fermentation is the foaming that occurs during the
manufacture of wine and beer, a process at least 10,000 years old. The frothing results
from the evolution of carbon dioxide gas, though this was not recognized until the 17th
century. French chemist and microbiologist Louis Pasteur in the 19th century used the
term fermentation in a narrow sense to describe the changes brought about
by yeasts and other microorganisms growing in the absence of air (anaerobically); he
also recognized that ethyl alcohol and carbon dioxide are not the only products of
fermentation.
Anaerobic breakdown of molecules
In the 1920s it was discovered that, in the absence of air, extracts of muscle catalyze the
formation of lactate from glucose and that the same intermediate compounds formed in
the fermentation of grain are produced by muscle. An important generalization thus
emerged: that fermentation reactions are not peculiar to the action of yeast but also
occur in many other instances of glucose utilization.

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Types of Chemical Reactions

glycolysis
The generation of pyruvate through the process of glycolysis is the first step in fermentation.
Glycolysis, the breakdown of sugar, was originally defined about 1930 as
the metabolism of sugar into lactate. It can be further defined as that form of
fermentation, characteristic of cells in general, in which the six-carbon sugar glucose is
broken down into two molecules of the three-carbon organic acid, pyruvic acid (the
nonionized form of pyruvate), coupled with the transfer of chemical energy to
the synthesis of adenosine triphosphate (ATP). The pyruvate may then be oxidized, in
the presence of oxygen, through the tricarboxylic acid cycle, or in the absence of oxygen,
be reduced to lactic acid, alcohol, or other products. The sequence from glucose to
pyruvate is often called the Embden–Meyerhof pathway, named after two German
biochemists who in the late 1920s and ’30s postulated and analyzed experimentally the
critical steps in that series of reactions.

The term fermentation now denotes the enzyme-catalyzed, energy-yielding pathway in

cells involving the anaerobic breakdown of molecules such as glucose. In most cells the
enzymes occur in the soluble portion of the cytoplasm. The reactions leading to the
formation of ATP and pyruvate thus are common to sugar transformation in muscle,
yeasts, some bacteria, and plants.
Industrial fermentation
Industrial fermentation processes begin with suitable microorganisms and specified
conditions, such as careful adjustment of nutrient concentration. The products are of
many types: alcohol, glycerol, and carbon dioxide from yeast fermentation of various
sugars; butyl alcohol, acetone, lactic acid, monosodium glutamate, and acetic acid from
various bacteria; and citric acid, gluconic acid, and small amounts of antibiotics, vitamin
B12, and riboflavin (vitamin B2) from mold fermentation. Ethyl alcohol produced via the
fermentation of starch or sugar is an important source of liquid biofuel.
The Editors of Encyclopaedia BritannicaThis article was most recently revised and updated by Adam
Augustyn.
tartaric acid
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tartaric acid
chemical compound
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Also known as: dihydroxybutanedioic acid
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William H. Brown
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tartaric acid
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Also called:

dihydroxybutanedioic acid
Key People:

Louis Pasteur
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polycarboxylic acid

racemic acid
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Tartaric acid, a dicarboxylic acid, one of the most widely distributed of plant acids,
with a number of food and industrial uses. Along with several of its salts, cream of tartar
(potassium hydrogen tartrate) and Rochelle salt (potassium sodium tartrate), it is
obtained from by-products of wine fermentation. In a partially purified form, tartar was
known to the ancient Greeks and Romans; the free acid was first isolated in 1769 by
Swedish chemist Carl Wilhelm Scheele. The lees, or sediments, and other waste
products from fermentation are heated and neutralized with calcium hydroxide; the
precipitated calcium tartrate is then treated with sulfuric acid to produce free tartaric
acid. Rochelle salt is prepared from the crude crystalline potassium acid salt, called
argol, by neutralization with sodium carbonate. Purified cream of tartar comes chiefly
from the filtrates from production of the acid and Rochelle salt. A third salt, tartar
emetic (antimony potassium tartrate), is made from the potassium acid salt and
antimony oxide.
Three stereoisomeric forms of tartaric acid exist: (1) dextrorotatory tartaric acid ( D-
tartaric acid) found in grapes and several other fruits, (2) levorotatory tartaric acid (L-
tartaric acid) obtained chiefly by resolution of racemic tartaric acid, and (3) a meso or
achiral form. Racemic tartaric acid (an equal mixture of D- and L-tartaric acid) is
prepared commercially by the molybdenum- or tungsten-catalyzed oxidation of maleic

anhydride with hydrogen peroxide.

Study of the crystallographic, chemical, and optical properties of the tartaric acids by
French chemist and microbiologist Louis Pasteur laid the basis for modern ideas
of stereoisomerism.

The various tartaric acids and the common tartrate salts are all colourless, crystalline
solids readily soluble in water. Tartaric acid is widely used as an acidulant in carbonated
drinks, effervescent tablets, gelatin desserts, and fruit jellies. It has many industrial
applications—e.g., in cleaning and polishing metals, in calico printing, in wool dyeing,
and in certain photographic printing and development processes. Rochelle salt is used in
silvering mirrors, in processing cheese, and in compounding mild cathartics. Cream of
tartar is incorporated into baking powders, hard candies, and taffies; and it is employed
in the cleaning of brass, the electrolytic tinning of iron and steel, and the coating of
other metals with gold and silver. Tartar emetic is used as an insecticide and a dyeing
mordant.
William H. Brown
rabies
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rabies
pathology
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Also known as: hydrophobia, lyssa
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Also called:

hydrophobia or lyssa
Key People:

Louis Pasteur

Sir Victor Horsley

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dog

disease

notifiable disease

rabies virus
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Rabies, acute, ordinarily fatal, viral disease of the central nervous system that is usually
spread among domestic dogs and wild carnivorous animals by a bite. All warm-blooded
animals, including humans, are susceptible to rabies infection. The virus, a rhabdovirus,
is often present in the salivary glands of rabid animals and is excreted in the saliva; thus,
the bite of the infected animal introduces the virus into a fresh wound. Under favourable
conditions, the virus propagates along nerve tissue from the wound to the brain and
becomes established in the central nervous system. After a time it spreads via nerves to
the salivary glands, where it frequently produces a foaming at the mouth.
The disease develops most often between four and six weeks after infection, but the
incubation period may vary from 10 days to eight months.

Rabies virus travels quickly in a bitten animal (e.g., raccoons, skunks, bats, foxes, dogs,
and cats, among other smaller animals) from the bite to the central nervous system. The
disease often begins with excitation of the central nervous system expressed as
irritability and viciousness. A rabid animal is most dangerous during the early stages of
the disease because it appears to be healthy and may seem friendly but will bite at the
slightest provocation. Wild animals that appear to be tame and that approach people or
human habitations in the daytime should be suspected of having rabies.

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Infected dogs usually show a short excitation phase that is characterized by restlessness,
nervousness, irritability, and viciousness and is followed by depression and paralysis.
After a few days they are unable to bite any more because the muscles of the throat are
paralyzed; they seek only a quiet place to hide and die from the rapid spread of
paralysis. Sudden death without recognizable signs of illness is also not uncommon.
Dogs that develop the predominantly excited type of rabies invariably die of the
infection, usually within three to five days after the onset of symptoms. Those that
develop the paralytic type of rabies without any evidence of excitation or viciousness
may recover on rare occasions. Paralysis of the “voice” muscles in rabid dogs may
produce a characteristic change in the sound of the bark.

Rabies in humans is similar to that in animals. Symptoms

include depression, headache, nausea, seizures, anorexia, muscle stiffness, and
increased production of saliva. Abnormal sensations, such as itching, around the site of
exposure are a common early symptom. The muscles of the throat become paralyzed so
that the person cannot swallow or drink, and this leads to a dread of water
(hydrophobia). The mental state of a person infected with rabies varies
from maniacal excitement to dull apathy—the term rabies means “madness”—but soon
the person falls into a coma and usually dies in less than one week owing to cardiac or
respiratory failure. Sometimes rabies is characterized by paralysis without any evidence
of excitation of the nervous system. In such cases the course of the disease may be
prolonged to a week or more.

There is no cure for rabies. The incubation period (the time that elapses between the
bite and the first symptom) is usually one to three months but in rare cases has been as
long as several years. This provides a chance to interrupt the otherwise inevitable
progress of the infection. The bite should be washed immediately because much, if not
all, of the virus can be thus removed. The bitten patient should then receive a dose
of antirabies serum. Serum is derived from horses or humans that have been immunized
with attenuated rabies virus; it provides the patient with already prepared antibodies
against the rabies antigen. The treatment is effective if given within 24 hours after
exposure but has little, if any, value if given three or more days after infection by rabies.

Active immunization with rabies vaccine should also be initiated to allow the patient’s
body to make its own antibody. The safest and most effective vaccines are human
diploid cell vaccine (HDCV), purified chick embryo cell culture (PCEC), and rabies
vaccine adsorbed (RVA). With older vaccines, at least 16 injections were required,
whereas with HDCV, PCEC, or RVA, 5 are usually sufficient. Persons at risk of rabies by
virtue of occupation (e.g., veterinarians) or travel to endemic areas should receive rabies
vaccine as a form of preexposure prophylaxis.
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The Editors of Encyclopaedia BritannicaThis article was most recently revised and updated by Adam
Augustyn.
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anthrax
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Also known as: malignant pustule, splenic fever, woolsorters’ disease
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Also called:

malignant pustule or woolsorters’ disease

Key People:

Louis Pasteur

Robert Koch
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biological weapon

zoonotic disease

notifiable disease

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 Bacillus anthracis
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Anthrax, acute, infectious, febrile disease of animals and humans caused by Bacillus
anthracis, a bacterium that under certain conditions forms highly resistant spores
capable of persisting and retaining their virulence for many years. Although anthrax
most commonly affects grazing animals such as cattle, sheep, goats, horses, and mules,
humans can develop the disease by eating the meat or handling the wool, hair, hides,
bones, or carcasses of affected animals. When anthrax—its name derived from the Greek
word for coal—attacks a person’s skin, a sore with a coal-black centre develops. Anthrax
spores can also be produced inexpensively and converted into either a powder or a
liquid, allowing anthrax to be used in a variety of weapons systems. Its potential use as a
biological weapon has made anthrax a prime concern of countries seeking to control the
spread of terrorism.
Anthrax in humans and animals
Anthrax in humans occurs as a cutaneous, pulmonary, or intestinal infection. The most
common type, cutaneous anthrax, occurs as a primary localized infection of the skin in
the form of a carbuncle. It usually results from handling infected material, lesions
occurring mostly on the hands, arms, or neck as a small pimple that develops rapidly
into a large vesicle with a black necrotic centre (the malignant pustule). There are
also bouts of shivering and chills, but there is little other disability. In more than 90
percent of the cases of anthrax in humans, the bacilli remain within the skin sore.
However, the bacilli may escape from the sore and spread via a lymph channel to the
nearest lymph node, where their spread is usually halted. Only seldom do the bacilli
invade the bloodstream, causing rapidly fatal septicemia (blood poisoning), internal
bleeding, and, sometimes, meningitis. The pulmonary form, called inhalation anthrax
(woolsorters’ disease), affects principally the lungs and pleura and results from inhaling
anthrax spores (e.g., in areas where hair and wool are processed). Inhalation anthrax is
occasionally transmitted to humans by spore-contaminated brushes or by wearing
apparel such as furs and leather goods. This form of the disease usually runs a rapid
course and terminates fatally due to the suffocating pneumonia that results. The
intestinal form of the disease, which sometimes follows the consumption of
contaminated meat, is characterized by an acute inflammation of the intestinal tract,
vomiting, and severe diarrhea.

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In all forms of anthrax, prompt diagnosis and early treatment are of great importance.
Antianthrax serum, arsenicals, and antibiotics (e.g., ciprofloxacin) are used with
excellent results. The hazard of infection to industrial workers can be reduced by
sterilization of potentially contaminated material before handling, wearing of protective
clothing, use of respirators, and sanitizing of facilities. Agricultural workers can be
safeguarded by vaccination and by avoiding the skinning or opening of animals that died
of the disease.

Farm animals that graze on contaminated pastures are susceptible to anthrax.

Outbreaks in swine, dogs, cats, and wild animals held in captivity generally result from
consumption of contaminated food. In the acute forms there is a rise in body
temperature followed by spasms, respiratory or cardiac distress, trembling, staggering,
and convulsions. Bloody discharges sometimes come from the natural body openings,
and edematous (serous fluid) swellings may appear on different parts of the body. Death
usually occurs within a day or two. Chronic anthrax occurs mostly in swine and dogs and
is characterized by marked swelling of the throat, difficult breathing, and a bloodstained
frothy discharge from the mouth. Affected animals sometimes die of suffocation.
Prophylactic vaccination is extensively used in preventing anthrax in livestock. During
outbreaks, strict quarantine measures, disposal of diseased carcasses by burning, fly
control, and good sanitation are essential in controlling the disease.

Anthrax is one of the oldest recorded diseases, being mentioned in the biblical book of
Exodus and among the Classical authors of Greek and Roman antiquity.
Devastating epidemics of the disease were recorded by many medieval and modern
writers. In the 16th to 18th century it sometimes spread across the southern part of
Europe, taking a heavy toll on human and animal life. The causative agent was identified
by French biologist Casimir-Joseph Davaine in 1863 and by German
bacteriologist Robert Koch, who isolated the organism in pure culture in 1876. An
effective vaccine was demonstrated by French chemist and microbiologist Louis
Pasteur in 1881. These discoveries were part of the origin and development of the
modern sciences of bacteriology and immunology.
Anthrax as a biological weapon
The bacterium that causes anthrax has a number of attributes that, in combination,
make it suitable as a biological weapon. In addition to being widely available—located
around the world in soil and in diseased animals and their remains—B. anthracis spores
are small enough to lodge readily in the lungs of humans. The anthrax bacterium has a
short incubation period and is highly lethal, requiring only a small amount to cause a
mass casualty effect. Indeed, aerosolized forms of anthrax sprayed over a large
population centre or a massed military force are capable of lethality approaching or
exceeding that of a nuclear weapon. Moreover, anthrax can be produced inexpensively,
in larger quantities than can other biological warfare agents, and in facilities that are
relatively easy to hide. It is more resilient to degradation from ultraviolet light than
most other biological agents. Finally, anthrax can be converted into either a powder or a
liquid, allowing it to be used in a number of types of weapons systems that utilize a
variety of delivery means, including missiles, bombers, artillery, mortars, or crop
dusters and similar aircraft.

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Anthrax has been weaponized by a number of states. Before it terminated its offensive
biological weapons program in 1969, the United States had a significant anthrax
weapons program. The former Soviet Union developed the world’s largest biological
weapons program, which it clandestinely continued 20 years beyond the date when it
signed the Biological Weapons Convention of 1972, which prohibited the development
and stockpiling of biological weapons. Iraq, under the rule of Ṣ addām Ḥ ussein, also
developed anthrax and a number of other biological weapons agents but claimed to have
destroyed such weapons after the Persian Gulf War of 1990–91. At least 14 other nations
are believed to have biological weapons programs.

Terrorists have used anthrax in an attempt to kill and frighten victims in both Japan and
the United States. The AUM Shinrikyo religious sect released anthrax in Tokyo on three
separate occasions in 1993, targeting downtown crowds and members of the Japanese
legislature. In 2001 a number of anthrax-laced letters were sent through the mail to the
offices of two U.S. senators and various media headquarters in New York and Florida,
killing five people along the letters’ routes and infecting more than a dozen others.

Several effective vaccines have been developed to protect against possible anthrax
infection, including Anthrax Vaccine Adsorbed (AVA), the vaccine developed to protect
United States military personnel. The anthrax vaccine can provide protection to most
recipients, although a small percentage do not acquire complete immunity. However, if
vaccinated military personnel were to encounter a massive dose of anthrax, such as
might be encountered on a battlefield, even a sensitized immune system can be
overwhelmed; a well-fitting mask with fine-grain filters is necessary to provide
protection in such instances.
The Editors of Encyclopaedia BritannicaThis article was most recently revised and updated by Adam
Augustyn.
immunization
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immunization
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Key People:

Clara Maass
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variolation

toxoid

inoculation
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Top Questions
What is immunization?
How does immunization work?
What is the difference between immunization and vaccination?
Immunization, process by which resistance to disease is acquired or induced in plants
and animals. This discussion focuses on immunization against infectious diseases in
vertebrate animals, specifically humans.

Immunization may occur naturally, as when a person is exposed unintentionally to a

pathogen (any infectious agent), or it may be brought about artificially through
a vaccine. In either case, immunization provides resistance, or immunity, to a particular
pathogen by means of antibody proteins that are targeted to eliminate that pathogen
from the body. These antibodies do not react to the entire pathogen but only to a specific
part of it, which is called an antigen. An individual can acquire immunity for a specific
pathogen passively or actively. In passive immunization a person receives antibodies
or lymphocytes that have been produced by another individual’s immune system; in
active immunization the individual’s own immune system is stimulated to produce
antibodies and lymphocytes.

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immune system: Prophylactic immunization

Passive immunization imparts immediate, but not long-lasting, protection against a

pathogen and may arise naturally, such as when a fetus receives antibodies from the
mother across the placenta or when a breast-feeding infant ingests antibodies in the
mother’s milk. Passive immunization against a particular pathogen, such as the hepatitis
B virus (HBV), also can be conferred artificially. A person lacking immunity to HBV can
receive a preparation called immune serum globulin that contains antibodies formed
against the virus. These antibodies are obtained from serum taken from animal or
human donors who previously were infected by or immunized against the virus.

Active immunization stimulates the immune system to produce antibodies against a

particular infectious agent. Active immunity can arise naturally, as when someone is
exposed to a pathogen. For example, an individual who recovers from a first case of
the measles is immune to further infection by the measles-causing virus, because the
virus stimulates the immune system to produce antibodies that specifically recognize
and neutralize the pathogen the next time it is encountered. Active immunization also
can be artificially induced through vaccination. Vaccines are preparations containing
antigens that stimulate an immune response without causing illness. The purpose of
vaccination is to ensure that a large enough number of antibodies and lymphocytes
capable of reacting against a specific pathogen or toxin are available before exposure to
it occurs. Active immunization is often long-lasting and may be reactivated quickly by a
recurrence of the infection or by revaccination.
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bacterial disease
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bacterial disease
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Bacterial disease, any of a variety of illnesses caused by bacteria. Until the mid-20th
century, bacterial pneumonia was probably the leading cause of death among the
elderly. Improved sanitation, vaccines, and antibiotics have all decreased the mortality
rates from bacterial infections, though antibiotic-resistant strains have caused a
resurgence in some illnesses. In the early 21st century, tuberculosis, which is caused
by Mycobacterium tuberculosis—several strains of which had developed resistance to
one or more drugs widely used to treat the infection—was among the
deadliest infectious diseases worldwide.

Bacteria cause disease by secreting or excreting toxins (as in botulism), by producing

toxins internally, which are released when the bacteria disintegrate (as in typhoid), or by
inducing sensitivity to their antigenic properties (as in tuberculosis). Other serious
bacterial diseases include cholera, diphtheria, bacterial meningitis, tetanus, Lyme
disease, gonorrhea, and syphilis.

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This article was most recently revised and updated by Kara Rogers.
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pasteurization
heating process
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Louis Pasteur
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ultra-high-temperature pasteurization

high-temperature-short-time method
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Pasteurization, heat-treatment process that destroys pathogenic microorganisms in
certain foods and beverages. It is named for the French scientist Louis Pasteur, who in
the 1860s demonstrated that abnormal fermentation of wine and beer could be
prevented by heating the beverages to about 57 °C (135 °F) for a few minutes.
Pasteurization of milk, widely practiced in several countries, notably the United States,
requires temperatures of about 63 °C (145 °F) maintained for 30 minutes or,
alternatively, heating to a higher temperature, 72 °C (162 °F), and holding for 15 seconds
(and yet higher temperatures for shorter periods of time). The times and temperatures
are those determined to be necessary to destroy Mycobacterium tuberculosis and other,
more heat-resistant, non-spore-forming, disease-causing microorganisms found in milk.
The treatment also destroys most of the microorganisms that cause spoilage and so
prolongs the storage time of food.

Ultra-high-temperature (UHT) pasteurization involves heating milk or cream to 138–

150 °C (280–302 °F) for one or two seconds. Packaged in sterile, hermetically sealed
containers, UHT milk may be stored without refrigeration for
months. Ultrapasteurized milk and cream are heated to at least 138 °C for at least two
seconds, but, because of less stringent packaging, they must be refrigerated. Shelf life is
extended to 60–90 days. After opening, spoilage times for both UHT and
ultrapasteurized products are similar to those of conventionally pasteurized products.

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Pasteurization of some solid foods involves a mild heat treatment, the exact definition of
which depends on the food. Radiation pasteurization refers to the application of small
amounts of beta or gamma rays to foods to increase their storage time.
The Editors of Encyclopaedia BritannicaThis article was most recently revised and updated by Kara
Rogers.