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Shinya Inoue: A Long Career Advancing Microscopy and Cell Biology (1921-2019).

Listen, observe, and be kind. These words characterize the great scientist and thoughtful human being that Shinya Inoue was. Following his passing on September 30, 2019, at the age of 98, tributes and remembrances poured in from around the world to the Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts, which has been Dr. Inoue's scientific home for over 30 years. His approach to scientific experimentation, technology development, kind teaching, and collegial interactions serve as a great example to us all. Since his passing, friends and colleagues emphasized the generosity, kindness, and patience that Dr. Inoue exhibited, along with his scientific rigor and fastidious and precise nature. This combination of traits is quite rare and clearly cherished by me and by all who were fortunate to be able to learn from him. I was lucky enough to experience one of his lectures at MBL during the Analytical and Quantitative Light Microscopy course (when he was a spry 87). Interactions with Dr. Inoue could be tough, as he was known to set high goals for himself and others, but with his gentle guidance and thoughtful feedback many of those goals were reached. His scientific achievements in live cell imaging were pioneering, demonstrating the existence of mitotic spindle fibers and chromosome movement, using his development of imaging technology; and his open sharing of ideas and results set an example and contributed to the progress in cell biology and cell motility.

Shinya Inoue was born in 1921 in London, England, the son of a diplomat, and grew up while his father was stationed abroad in several different countries. He returned to Japan for high school and his higher education. In his first year at Tokyo University, Inoue met Professor Katsuma Dan, who sparked his interest in experimental science. Dr. Dan had an approach to teaching that was more investigative than was typical at the time in Japanese schools. Later, Prof. Dan challenged Inoue to build a microscope that would allow the observation of the dynamic cellular structure of the mitotic spindle in the sea urchin embryo. Inoue's work at that time displayed the ingenuity, perseverance, and work ethic that would continue throughout his life. He built his first polarized light microscope from a discarded machine gun base and a tin tea can--parts found in the area of the Misaki Marine Biological Station, Japan, just after WWII. This achievement was later featured in Ripley's Believe It or Not! Subsequently, he built and improved upon at least seven "Shinya-scopes." His most recent development was a centrifugation polarizing microscope that can exert forces on cellular components of different densities inside living cells, including their stratification. To freeze the fast rotor motion and allow stroboscopic imaging, the specimen is illuminated with nanosecond laser pulses that are synchronized to the rotor position. Inoue's collaborations with many commercial partners were key to his hardware developments.

Shinya Inoue arrived at Princeton in 1948 with the help of a fellowship that scientist Jean Dan (wife of Katsuma Dan) helped secure. He was able to travel to the Marine Biological Laboratory during his first summer in the United States. Inoue returned to Woods Hole to perform research each summer throughout his time as a student (PhD, Princeton 1951) and then as faculty at Dartmouth College (1959-1966) and the University of Pennsylvania (Professor, 1966-1982). He became a permanent Marine Biological Laboratory Senior Scientist in 1982, an MBL Distinguished Scientist in 1986, and founder of the Architectural Dynamics in Living Cells Program in 1992. As a collaborative effort of commercial and academic representatives, he founded an intensive light microscopy course at MBL, where he served as course director (1979-1987) and lecturer for many years. In 1998 he was invited to join the Editorial Board of The Biological Bulletin to help develop and edit its series on "Concepts of Imaging and Microscopy." A number of articles were published over the next several years that continued to emphasize the collaboration between academic and commercial scientists, in order to educate users of those instruments about the rapid advances in the field. Inoue continued as Section Editor and Associate Editor on this journal's Editorial Board from 1998 until his death.

He is probably best known for two very important and pioneering achievements: he showed that mitotic spindle fibers were responsible for segregating chromosomes during mitosis, and he developed light microscopy techniques for viewing cell motility and cytoskeleton in live cells. His technique improved polarized light microscopy, allowing the visualization of internal structures of living cells in order to determine the mechanism of cell division and the major structural protein components responsible. Standard bright-field or even phase-contrast microscopy was not able to show detailed information about the fibers that had been proposed to be part of the mitotic spindle. Using his custom-built polarizing microscope, Inoue clearly demonstrated the reality of spindle fibers, now known as microtubules, responsible for chromosome movements (Inoue, 1953. Chromosoma 5: 487-500). He postulated that growing and shortening of mitotic spindle fibers could generate forces to move chromosomes, leading to the discovery of a whole new type of motility different from myosin muscle filament sliding (Inoue and Sato, 1967. J. Gen. Physiol. 50: 259-292). Polymerization and depolymerization of a biological polymer drive chromosome movement, and Inoue and Sato's suggestion that this motility could move other organelles or push on biological membranes was recognized as a major breakthrough in cell biology (Grisendi, 2010. Nat. Rev. Mol. Cell Biol. 9: s8-s9).

Over the years Inoue's improvements in microscopy continued: he was also the co-inventor of video microscopy and spurred more extensive use of live cell imaging in biology. Using a video camera to record images (Inoue, 1981. J. Cell Biol. 89: 346-356; also independently by Robert Allen) with computer-assisted contrast enhancement, he was able to visualize structures inside cells with greater clarity and detail than had been previously accomplished.

His successful approach of developing new technologies in microscopy in order to answer critical questions in (cell) biology was the key to his contributions to science and teaching. Many scientists have emphasized this about their interactions with him. Shinya would ask, "What is your biological question?" Then he would expect you to figure out how to answer your question and develop microscopy (or other) tools needed to do so. He knew that technology development should be rooted in the pursuit of answering a biological question, rather than building new equipment for the sake of inventing.

Many people appreciated Inoue's teaching, mentorship, and collegiality. His colleagues describe him as both exacting and demanding while also being patient, generous, and helpful. He had a dedication to his work that often stretched into the late-night hours. And he was, in fact, known to prefer working later (and starting later). One collaborator described a rewarding grant-writing experience with Inoue, who would send suggestions and comments past midnight when he was over 90 years old. Inoue's drive to help everyone he encountered to improve is what is most revered by his colleagues. He taught many, many people in both the MBL Physiology course and the Microscopy course (now called AQLM: Analytical and Quantitative Light Microscopy), along with many individual and collaborative interactions.

Beyond his scientific generosity, Shinya Inoue was a significant philanthropist to the Marine Biological Laboratory and others. He supported his lab members financially at times when grants did not come through. He helped students and visitors arrange for housing in Woods Hole and often hosted visitors at his house. He contributed to friends' personal goals or needs, for example, by sponsoring a charity marathon run or by helping purchase solar panels or a furnace for a friend's house.

Shinya Inoue was recognized for his achievements with many awards, perhaps most notably by the International Prize for Biology from the Japan Society for the Promotion of Science (2003). He is also a member of the National Academy of Sciences and the American Academy of Arts and Sciences. In the summer of 2011, friends and colleagues held an honorary symposium for him. It was aptly titled "Listening to Life Through the Microscope."

Shinya Inoue has left a lasting legacy, not only of scientific discovery and technology development but also, more importantly, of personal kindness and generosity. As colleague Stephen Smith said in his written tribute, Inoue "has inspired us all to treat each other a little bit better as we go about our work." His sharing and openness in science should be an example to us all, and we can keep his legacy alive by both striving for new knowledge about the mechanisms of life and showing decency and graciousness to one another.

--Lisa A. Cameron

Editorial Board

The Biological Bulletin

DOI: 10.1086/707517
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Author:Cameron, Lisa A.
Publication:The Biological Bulletin
Date:Dec 1, 2019
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