Burnet, Frank Macfarlane WORLD OF MICROBIOLOGY AND IMMUNOLOGY
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buffer system compensates for such additions and maintains
the pH within the required range.
This buffering system is intimately tied to respiration,
and an exceptional feature of pH control by this system is the
role of ordinary breathing in maintaining the pH. Carbon diox-
ide is a normal product of metabolism. It is transported to the
lungs, where it is eliminated from the body with every exhala-
tion. However, carbon dioxide in blood is converted to car-
bonic acid, which dissociates to produce the hydrogen
carbonate ion and the hydronium ion. If a chemical reaction or
the ingestion of an acidic material increases the hydronium ion
concentration in the blood, bicarbonate ion reacts with the
added hydronium ion and is transformed into carbonic acid.
As a result the concentration of dissolved carbon dioxide in
the blood increases. Respiration increases, and more carbon
dioxide is expelled from the lungs. Conversely, if a base is
ingested, the hydronium ion reacts with it, causing a decrease
in the concentration of hydronium ion. More carbonic acid
dissociates to restore the hydronium ion consumed by the
base. This requires more carbon dioxide to be dissolved in the
blood, so respiration is decreased and more gas is retained.
To act as a buffer, a solution must maintain a nearly con-
stant pH when either acid or base is added. Two considerations
must be made when a buffer is prepared: (1) Which pH is
desired to maintain? The desired pH defines the range of the
buffer. (2) How much acid or base does the solution need to
consume without a significant change in pH? This defines the
capacity of the buffer. The desired pH also determines the
compounds used in making up the buffer. The quantity of acid
or base the buffer must be able to consume determines the
concentrations of the components that must be used, and
which allows biological reactions to take place consistently.
See alsoBiochemical analysis techniques; Laboratory tech-
niques in microbiology
BURNET, FRANKMACFARLANE
(1899-1985)Burnet, Frank Macfarlane
Australian immunologist and virologist
While working at the University of Melbourne’s Walter and
Eliza Hall Institute for Medical Research in the 1920s, Frank
Macfarlane Burnet became interested in the study of viruses
and bacteriophage(viruses that attack bacteria). That interest
eventually led to two major and related accomplishments. The
first of these was the development of a method for cultivating
viruses in chicken embryos, an important technological step
forward in the science of virology. The second accomplish-
ment was the development of a theory that explains how an
organism’s body is able to distinguish between its own cells
and those of another organism. For this research, Burnet was
awarded a share of the 1960 Nobel Prize for physiology or
medicine (with Peter Brian Medawar).
Burnet was born in Traralgon, Victoria, Australia. His
father was Frank Burnet, manager of the local bank in
Traralgon, and his mother was the former Hadassah Pollock
MacKay. As a child, Burnet developed an interest in nature,
particularly in birds, butterflies, and beetles. He carried over
that interest when he entered Geelong College in Geelong,
Victoria, where he majored in biology and medicine.
In 1917, Burnet continued his education at Ormond
College of the University of Melbourne, from which he
received his bachelor of science degree in 1922 and then, a
year later, his M.D. degree. Burnet then took concurrent posi-
tions as resident pathologist at the Royal Melbourne Hospital
and as researcher at the University of Melbourne’s Hall
Institute for Medical Research. In 1926, Burnet received a
Beit fellowship that permitted him to spend a year in residence
at the Lister Institute of Preventive Medicine in London. The
work on viruses and bacteriophage that he carried out at Lister
also earned him a Ph.D. from the University of London in
- At the conclusion of his studies in England in 1928,
Burnet returned to Australia, where he became assistant direc-
tor of the Hall Institute. He maintained his association with the
institute for the next thirty-seven years, becoming director
there in 1944. In the same year, he was appointed professor of
experimental medicine at the University of Melbourne.
Burnet’s early research covered a somewhat diverse
variety of topics in virology. For example, he worked on the
classification of viruses and bacteriophage, on the occurrence
of psittacosis in Australian parrots, and on the epidemiologyof
herpesand poliomyelitis. His first major contribution to virol-
ogy came, however, during his year as a Rockefeller fellow at
London’s National Institute for Medical Research from 1932
to 1933. There he developed a method for cultivating viruses
in chicken embryos. The Burnet technique was an important
breakthrough for virologists since viruses had been notori-
ously difficult to cultureand maintain in the laboratory.
Over time, Burnet’s work on viruses and bacteriophage
led him to a different, but related, field of research, the verte-
brate immune system. The fundamental question he attacked
is one that had troubled biologists for years: how an organ-
ism’s body can tell the difference between “self” and “not-
self.” An organism’s immune system is a crucial part of its
internal hardware. It provides a mechanism for fighting off
invasions by potentially harmful—and sometimes fatal—for-
eign organisms (antigens) such as bacteria, viruses, and fungi.
The immune system is so efficient that it even recognizes and
fights back against harmless invaders such as pollen and dust,
resulting in allergic reactions.
Burnet was attracted to two aspects of the phenomenon
of immunity. First, he wondered how an organism’s body dis-
tinguishes between foreign invaders and components of its
own body, the “self” versus “not-self” problem. That distinc-
tion is obviously critical, since if the body fails to recognize
that difference, it may begin to attack its own cells and actu-
ally destroy itself. This phenomenon does, in fact, occur in
some cases of autoimmune disorders.
The second question on which Burnet worked was how
the immune system develops. The question is complicated by
the fact that a healthy immune system is normally able to rec-
ognize and respond to an apparently endless variety of anti-
gens, producing a specific chemical (antibody) to combat each
antigenit encounters. According to one theory, these antibod-
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