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Bud Ryan
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Bud
Ryan
Clarence A.
“Bud” Ryan, Charlotte Y. Martin Professor of Biochemistry
and a Fellow of the Institute of Biological Chemistry, Washington
State University (WSU), died on October 7, 2007, at the age of 76.
Bud was widely known for his work on the regulation and function
of plant proteinase inhibitors in defense against insect herbivores.
In addition to being an excellent scientist, Bud was a cherished
mentor to dozens of students and postdocs, and he was dedicated
to promoting plant biology at the university, national, and international
levels.
Bud was born
on September 29, 1931, in Butte, Montana, the second of four children.
When Bud was 10 years old, the family moved to Helena, Montana.
Bud became interested in chemistry at Cathedral High School and,
after graduation, decided to enroll at Carroll College in Helena.
He paid his way through college by cleaning a bar and working as
a ticket taker at the local movie theater. As a starter for the
conference-winning basketball team for two years, Bud would maintain
his passion for the game—as both a participant and a spectator—for
the rest of his life. Bud received his bachelor’s degree in
chemistry with a minor in bacteriology from Carroll College in June
1953.
After college,
Bud worked for the Montana State Highway Laboratory and as a cabbie
at night to pay off his college loans. During this time, he met
Patricia Meunier, the love of his life, at a local barn dance. Bud
and Pat were married the following May, in 1954.
Bud entered
the graduate program at Montana State University in 1954. He conducted
his thesis research, entitled “A New Transglucosidase Found
in Potatoes,” in the laboratory of Dr. Kenneth Goering, and
in June 1959 he obtained his PhD in chemistry. After graduation,
Bud received several job offers to work in industry. With a young
family (Bud and Pat’s children, Jamie, Steven, and Janice,
were born while Bud was in graduate school), a career in industry
seemed to be the best route for the Ryan family.
But by this
time Bud had been bitten by the research bug. Relying on intuition,
as he did so often during his life, Bud decided to pursue an opportunity
to do postdoctoral work. He first studied with Dr. T. E. King at
Oregon State University and then at the USDA Western Regional Laboratory
in Albany, California, with enzymologist Dr. A. K. Balls. It was
in Albany that Bud came across a research article describing how
potato peels inhibit cholinesterase activity. Bud wondered what
the plant chemical might be and whether it would inhibit the esterase
activity of trypsin and chymotrypsin.
Always quick
to improvise, Bud ran down to the grocery store and purchased a
bag of potatoes and, within six months, had crystallized the first
chymotrypsin inhibitor from plants. Based on this discovery, Bud
received a Career Development Award from the National Institutes
of Health to further study the properties of the inhibitor and to
purify other inhibitors. The course for Bud’s long and productive
career in plant biology was set.
Bud and his
family moved in 1964 to Pullman, Washington, where he accepted an
assistant professor position in the Department of Agricultural Chemistry
at WSU. In 1966 a fourth child, Joseph Patrick (Joe Pat) Ryan, was
born. Bud would spend the rest of his career at WSU. His decision
to remain at WSU was influenced by the stimulating research environment
in the Department of Agricultural Chemistry, which, in 1980, became
the Institute of Biological Chemistry (IBC). Throughout his career,
Bud played a prominent role in shaping the Institute into a world-renowned
research facility.
Bud’s early
success in characterizing the biochemical properties of proteinase
inhibitors (PIs) led him to investigate factors that control their
accumulation in plants. He discovered, for example, that chymotrypsin
inhibitor I accumulates transiently in potato leaves as a temporary
storage protein before protein resources are allocated to the developing
tubers. Extending his studies on PIs to other plant species, Bud
observed the occasional and somewhat unpredictable accumulation
of PI-I in tomato leaves, and he suspected that the expression of
the inhibitor was influenced by environmental conditions.
On investigating
this phenomenon more closely, Bud and Terry Green (a postdoc in
the lab) found that tissue damage inflicted by Colorado potato beetles
resulted in massive accumulation of PIs in potato and tomato leaves.
This landmark discovery, which was published in Science in 1972,
suggested that wound-inducible PI expression makes the host plant
less nutritional and perhaps lethal to invading insects. The paradigm
that plants, rather than being passive victims of insect assault,
respond dynamically to herbivory through the production of defensive
compounds pervades much of the molecular plant–insect interaction
research to this day.
The 1972 Green
and Ryan Science paper also reported that wounding of a single leaf
causes PI expression in undamaged aerial tissues of the plant. This
discovery implied that plants possess an intercellular communication
system in which signals generated at the wound site are propagated
to distal leaves to warn of impending danger. Wound-induced systemic
expression of PIs has been adopted by many laboratories as a model
system for studying long-distance signaling in plants.
Tragically,
in 1976, Bud and Pat’s oldest son, Steven, died in a scuba
diving class accident. Later the same year, Joe Pat was killed when
the car he was riding in was struck by a drunk driver. Somehow,
Bud and Pat persevered through these terrible events.
By the late
1980s, Bud was focusing his research on understanding the signal
transduction pathway leading to PI gene expression in wounded tomato
leaves. But where should one start in such a project? Brady Vick
and Don Zimmerman, researchers at the USDA Agricultural Research
Service in Fargo, North Dakota, had recently elucidated the biosynthetic
route of an interesting linolenic acid derivative called jasmonic
acid (JA). At that time, JA’s function was unknown, but experiments
being conducted in Japan, Germany, and the United States suggested
that it had various biological activities, including stimulation
of the accumulation of “jasmonate-inducible proteins”
of unknown function.
Fortunately,
Bud had a nearly photographic memory of previous experiments conducted
in the lab, and he recalled an early unpublished experiment by Mary
Kay Walker-Simmons showing that linolenic acid treatment could stimulate
PI production. Maybe linolenic acid was metabolized to JA to induce
PIs? The first sample of JA to come into Bud’s lab was found
on the WSU campus. The compound, in the form of methyl-JA (MeJA),
was sprayed onto tomato plants. The following day, the result was
spectacular: MeJA had captured the laboratory record for a compound
that induces the highest level of PI accumulation. It thus became
clear that JA is an important regulator of plant defense responses.
These early experiments performed in Bud’s lab played an important
role in transforming JA from a relatively obscure phytochemical
to a full-fledged member of the plant hormone family.
Interestingly,
the inducing effect of MeJA appeared to spread from the sprayed
plant to nearby control plants, presumably as a result of MeJA’s
volatility. This observation immediately raised the exciting possibility
that plants able to produce MeJA might be able to stimulate PI gene
expression in neighboring tomato plants. Indeed, this prediction
was confirmed in a now-classic Ryan lab experiment in which MeJA
emitted from sagebrush (Artemisia tridentata) induced PI expression
in nearby tomato plants. Although this experiment was conducted
in an artificially closed system, the unequivocal demonstration
of interplant communication sparked intense excitement within the
community of plant biologists and ecologists. The role of plant
volatiles in mediating plant-to-plant communication under natural
conditions remains a highly active research area to this day.
Well before
the discovery of JA and MeJA as endogenous regulators of PI expression,
Bud’s lab had been engaged in a long-term study to identify
PI-inducing factors from tomato leaves. This effort relied on a
simple bioassay in which fractionated leaf extracts were tested
for PI-inducing activity in an immunodiffusion assay. In 1991, after
tens of thousands of such bioassays, this work paid off with the
discovery of systemin, which was the first peptide signal identified
in plants. Bud’s group would go on to publish a series of influential
papers describing systemin’s role in the wound response of
tomato plants.
In 1999, at
the age of 68, Bud “retired” from university administrative
and teaching duties, only to take a more active role in his research
on plant peptide signaling. During this period, Bud and his colleagues
would discover several new classes of bioactive peptides, including
hydroxyproline-rich glycopeptides (HypSys), rapid alkalization factors
(RALFs) that regulate root development, and a small polypeptide
(Pep1) from Arabidopsis involved in the control of defense responses
against pathogen attack. As was the case for tomato systemin, these
novel peptides are derived from proteolytic processing of larger
precursor proteins. These pioneering contributions to our understanding
of peptide signaling in higher plants show that Bud remained fully
engaged at the cutting edge of plant biology research even after
“retirement.”
Bud received
numerous honors for his outstanding contributions to science, including
election to the U.S. National Academy of Sciences (1987) for his
early work on proteinase inhibitors and plant–herbivore interactions.
He was the first faculty member from WSU elected to the academy.
Bud received the Stephen Hales Prize from the American Society of
Plant Physiologists (1992) and was a member of both the academic
and athletic halls of fame at Carroll College. In recognition of
his outstanding and long-term contributions to plant biology and
service to the Society, Bud was named a member of the inaugural
class of ASPB Fellows in 2007.
To many of us
who were fortunate enough to work with Bud, he will always be remembered
as a wonderful mentor and colleague, whose enthusiasm for science
was contagious. With modesty and a steady stream of humor, Bud created
a stimulating research environment in which students and postdocs
could thrive. As the jasmonate and systemin stories were unfolding
in the late 1980s and through the 1990s, the work was often carried
out at a fast pace and with a sense of excitement and camaraderie.
The “hot” results of the day were typically discussed
around the coffee pot, and new experiments were planned. The atmosphere
that Bud created made his laboratory an exceptional place in which
to conduct research.
Bud was in good
health up until the time of his death. He was an avid golfer and
fisherman. Until the last two years, he was still playing basketball
with the noon group at the gym. When he finally retired from basketball,
he took up a new sport, ice skating. Proving it is never too late
to learn something new, Bud and Pat took skating lessons and hit
the ice before work twice a week.
Bud Ryan is
survived by his loving wife Pat, his two daughters, Jamie Ryan and
Janice Thrall (Terry), and two special granddaughters, Kymberly
and Haleigh Thrall. The family suggests that contributions be directed
to the Steve & Joe Pat Ryan Memorial Fund, Carroll College,
1601 N. Benton Ave., Helena MT 59625-0002.
Ted Farmer
University of Lausanne
Gregg Howe
Michigan State University
Greg Pearce
Washington State University
Andreas Schaller
University of Hohenheim
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