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Thad L. Beyle Distinguished Professor of Biology. Combined genetics, molecular biology and live-cell microscopy to illuminate understanding of the way centromeres and kinetochores function in the proper segregation of chromosomes. Was first to determine the structure of a eukaryotic centromere and to show it had a nucleosomal foundation. His lab developed a transcriptionally inactivated conditional centromere that opened new ways to study mechanisms that cause chromosome nondisjunction and aneuploidy and was first to visualize the budding yeast nucleus and spindle microtubule dynamics in live cells and quantify their dynamic changes throughout the cell cycle. Recent breakthrough was the application of quantitative and super-resolution fluorescence microscopy techniques to measure protein copy number within the centromere at the inner kinetochore and within the microtubule attachment site at the outer kinetochore. Conducted the first careful analysis of chromosome structure around centromeres, culminating in a 3-D model of centromeric DNA organization and revealing how mechanical properties of DNA contribute to fidelity of chromosome segregation by the spindle.