WORLD OF MICROBIOLOGY AND IMMUNOLOGY Kluyver, Albert Jan325•
nine tRNA. The nucleotide sequence of this tRNA had been
determined in Robert Holley’s laboratory. In 1970, when
Khorana announced the total synthesis of the first wholly arti-
ficial gene, his achievement was honored as a major landmark
in molecular biology. Six years later, Khorana and his associ-
ates synthesized another gene. In the 1980s, Khorana carried
out studies of the chemistry and molecular biology of the gene
for rhodopsin, a protein involved in vision.
In 1966, Khorana was elected to the National Academy
of Sciences. His many honors and awards include the Merck
Award from the Chemical Institute of Canada, the Dannie-
Heinneman Prize, the American Chemical Society Award for
Creative Work in Synthetic Organic Chemistry, the Lasker
Foundation Award for Basic Medical Research, the Padma
Vibhushan Presidential Award, the Ellis Island Medal of
Honor, the National Medal of Science, and the Paul Kayser
International Award of Merit in Retina Research. He holds
Honorary Degrees for numerous universities, including Simon
Fraser University, Vancouver, Canada; University of
Liverpool, England; University of Punjab, India; University of
Delhi, India; Calcutta University, India; University of Chicago;
and University of British Columbia, Vancouver, Canada.See alsoGenetic regulation of eukaryotic cells; Microbial
geneticsKKitasato, Shibasaburo ITASATO, SHIBASABURO(1852-1931)
Japanese bacteriologistBacteriologist Shibasaburo Kitasato made several important
contributions to the understanding of human disease and how
the body fights off infection. He also discovered the bacterium
that causes bubonic plague.
Born in Kumamoto, Japan, Kitasato, completed his med-
ical studies at the University of Tokyo in 1883. Shortly after, he
traveled to Berlin to work in the laboratory of Robert Koch.
Among his greatest accomplishments, Kitasato discovered a
way of growing a pure cultureof tetanusbacillus using anaer-
obic methods in 1889. In the following year, Kitasato and
German microbiologist Emil von Behringreported on the dis-
covery of tetanus and diphtheriaantitoxin. They found that ani-
mals injected with the microbes that cause tetanus or diphtheria
produced substances in their blood, called antitoxins, which
neutralized the toxins produced by the microbes. Furthermore,
these antitoxins could be injected into healthy animals, provid-
ing them with immunityto the microbes. This was a major find-
ing in explaining the workings of the immune system. Kitasato
went on to discover anthraxantitoxin as well.
In 1892, Kitasato returned to Tokyo and founded his
own laboratory. Seven years later, the laboratory was taken
over by the Japanese government, and Kitasato was appointed
its director. When the laboratory was consolidated with the
University of Tokyo, however, Kitasato resigned and founded
the Kitasato Institute.
During an outbreak of the bubonic plague in Hong
Kong in 1894, Kitasato was sent by the Japanese government
to research the disease. He isolated the bacterium that causedthe plague. (Alexandre Yersin, 1863 – 1943, independently
announced the discovery of the organism at the same time).
Four years later, Kitasato and his student Kigoshi Shiga were
able to isolate and describe the organism that caused one form
of dysentery.
Kitasato was named the first president of the Japanese
Medical Association in 1923, and was made a baron by the
Emperor in 1924. He died in Japan in 1931.See alsoAntibody and antigen; Bacteria and bacterial infec-
tion; Immunity, active, passive and delayed; ImmunizationKKluyver, Albert JanLUYVER, ALBERTJAN(1888-1956)
Dutch microbiologist, biochemist, and botanistAlbert Jan Kluyver developed the first general model of cell
metabolismin both aerobic and anaerobic microorganisms,
based on the transfer of hydrogen atoms. He was a major
exponent of the “Delft School” of classical microbiology in
the tradition of Antoni van Leeuwenhoek (1632–1723).
Outside Delft, he also drew on the legacy of Louis Pasteur
(1822–1895), Robert Koch (1843–1910), and Sergei
Nikolayevich Winogradsky (1856–1953).
Born in Breda, the Netherlands, on June 3, 1888,
Kluyver was the son of a mathematician and engineer, Jan
Cornelis Kluyver, and his wife, Marie, née Honingh. In 1910,
he received his bachelor’s degree in chemical engineering
from the Delft University of Technology, but immediately
shifted his focus toward botany and biochemistry, winning his
doctorate in 1914 with a dissertation on the determinations of
biochemical sugars under the tutelage of Gijsebertus van
Iterson, professor of microscopic anatomy. In 1916, on van
Iterson’s recommendation, the Dutch government appointed
Kluyver as an agricultural and biological consultant for the
Dutch East Indies colonial administration.
In 1921, again on van Iterson’s recommendation,
Kluyver succeeded Martinus Willem Beijerinck(1851–1931) as
director of the microbiology laboratory at Delft, where he
spent the rest of his career. He immediately acquired the most
modern equipment and established high standards for both
collegiality and research. The reorganized laboratory thrived.
Kluyver’s reputation soon attracted many excellent graduate
students, such as Cornelius Bernardus van Niel (1897–1985),
another chemical engineer. Van Niel received his doctorate
under Kluyver with a dissertation on propionic acid bacteriain
1928 and was immediately offered an appointment at Stanford
University.
In a landmark paper, “Eenheid en verscheidenheid in de
stofwisseling der microben” [Unity and diversity in the metab-
olism of microorganisms] Chemische Weekblad, Kluyver
examined the metabolic processes of oxidation and fermenta-
tionto conclude that, without bacteria and other microbes, all
life would be impossible. Two years later he co-authored with
his assistant, Hendrick Jean Louis Donker, another important
paper, “Die Einheit in der Biochemie” [Unity in biochemistry]
Chemie der Zelle und Gewebe, which asserted that all life
forms are chemically interdependent because of their sharedwomi_K 5/6/03 3:29 PM Page 325