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Proc Natl Acad Sci U S A
Proc Natl Acad Sci U S A
PNAS
Proceedings of the National Academy of Sciences of the United States of America
0027-8424
1091-6490
National Academy of Sciences

39196638
202413204
10.1073/pnas.2413204121
retroRetrospective97
Retrospective
William B. Wood: Pioneering scientist and educator (1938 to 2024)
Strome Susan sstrome@ucsc.edu
a 1 https://orcid.org/0000-0001-9496-7412

McIntosh J. Richard b
aDepartment of Molecular, Cell and Developmental Biology, University of California Santa Cruz, Santa Cruz, CA 95064
bDepartment of Molecular, Cellular and Developmental Biology, University of Colorado at Boulder, Boulder, CO 80309
1To whom correspondence may be addressed. Email: sstrome@ucsc.edu.
28 8 2024
17 9 2024
28 8 2024
121 38 e2413204121Copyright © 2024 the Author(s). Published by PNAS.
2024
https://creativecommons.org/licenses/by-nc-nd/4.0/ This article is distributed under Creative Commons Attribution-NonCommercial-NoDerivatives License 4.0 (CC BY-NC-ND).

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pmcWilliam Barry (Bill) Wood, Distinguished Professor Emeritus in the Department of Molecular, Cellular and Developmental Biology (MCDB) at the University of Colorado at Boulder, died on May 9, 2024, at age 86. Bill was an outstanding researcher, a pioneer in teaching innovation, and a gifted musician. He inspired a generation of young scientists and teachers with his insightful approaches to imaginative research and his intense focus on education. His ability to lead with humility, kindness, and empathy was exceptional. He will be missed by colleagues and friends at home and abroad.

Bill feeling cheerful after a good concert at the Frasier Retirement community, Boulder, CO, 2023. Image Credit: Alex Herzog (photographer).

The eldest of five children, Bill grew up in a home full of science, scholarship, and athletics. His mother had studied bacteriology and briefly taught high school science. His father, an all-American football player in college, was a physician and medical researcher, first at Washington University in St. Louis and later at Johns Hopkins Medical School in Baltimore. Coached by his father, Bill became an exceptional tennis player, winning the Maryland Junior State Tennis Championship at age 17 and going on to join Harvard’s varsity tennis team.

Bill had a lifelong passion for music. In high school, he sang and played guitar with fellow student and future bluegrass great John Hartford. As a Harvard undergraduate, he hosted “Balladeers,” a weekly program on the campus radio station, through which he met a young, unknown musician named Joan Baez. The two performed in Cambridge coffee houses and in 1959 made a record together called “Folksingers ‘Round Harvard Square.” The next year, Joan produced her smash solo album, “Joan Baez,” while Bill opted to pursue a career in science. In an interview conducted at Stanford in 2015, Wood reflected on that early fork in the road. He recalled with a modest smile: “It was very clear which of us should go to grad school and which should go to the Newport Folk Festival and become a star.” Pausing, he added: “I have to say, with all due respect to the scientists, that was the most fun I ever had, singing with Joan Baez.”

After Bill finished his degree in Chemistry at Harvard, he went west to Stanford and earned a PhD in Biochemistry with Nobel laureate Paul Berg. Following a brief postdoc with Werner Arber at the University of Geneva, Bill joined the Department of Biology at Caltech as an Assistant Professor; there he collaborated with his new colleague Bob Edgar.

The Wood and Edgar team undertook a novel scientific approach, combining biochemistry (Wood) and genetics (Edgar) to figure out the assembly pathway of the virus, bacteriophage T4. These bacteriophages (bacteria eaters), which look like lunar landing modules, are composed of a head containing the viral DNA, a tail tube through which the viral DNA is injected into bacterial cells, and tail fibers that mediate attachment to bacteria.

Bill (Right) and Bob Edgar (Left) at Caltech circa 1966 studying a T4 phage assembly test plate. Image Credit: Walt Girdner (photographer).

Edgar et al. had previously treated T4 phages with chemicals that damage DNA, making mutant phages that could still infect bacteria but failed to produce new infective phages. They assigned mutants to “complementation groups” by infecting bacteria simultaneously with two different strains of mutant phage. If the mutations were in different genes, the two viral genomes complemented one another, each providing a functional copy of the gene that was mutant in the other. The doubly infected bacteria produced infective phages.

Bill moved this “mixing different mutants” approach from inside living cells to working with cell lysates and achieved stunning results. When he broke open two populations of bacteria, each infected with a different T4 mutant, and mixed the lysates, the cell-free mixture could assemble infective phages. This process became known as “in vitro” (in glass) complementation, because the mutant phages complemented each other in a test tube without a living cell to help.

This innovation, along with electron microscopy, enabled Wood, Edgar, and others to determine which parts of phage T4 were encoded by each phage gene and to piece together a pathway for phage assembly from the protein products of the individual phage genes. This tour de force established a new paradigm for the assembly of complex biological structures and contributed to Bill’s election into the National Academy of Sciences at the young age of 34.

In vitro complementation became an important tool for many scientists studying diverse processes. Bruce Alberts noted that it formed the basis of his early work on DNA replication.

After Bill’s success with T4 phage assembly, he turned his attention to the more complex process of how one-cell embryos develop into multicellular adults with diverse tissues. He selected a model system newly established by Sydney Brenner, the nematode worm Caenorhabditis elegans. This tiny animal, only one millimeter long, can be fed on bacteria in the laboratory and grows to adulthood in just three days, making it easy, cheap, and fast to study. To learn worm biology, Bill did a sabbatical with David Hirsh in the Department of MCDB at the University of Colorado in (CU) Boulder.

Excited by his research experience in Boulder, Bill considered setting up a lab at Caltech to study worm development. At the time, however, Hirsh’s department was seeking a chair who could lead them in exciting new directions. Several enthusiastic members of the department flew to Pasadena and convinced Bill to move to Colorado as their new chair.

Bill’s lab at CU Boulder tackled several important developmental questions, ranging from how cell fates are determined in embryos to how worms are instructed to develop as hermaphrodites versus males.

In the cell-fate arena, experts were debating whether the developmental fates and division patterns of individual cells in early embryos are determined by internal factors that are packaged into those cells or determined by external cues from neighboring cells. Bill’s group found compelling evidence for both mechanisms, a finding that has been borne out across organisms.

Bill’s lab also studied an unusual type of organelle that is not bound by a membrane. They showed that these organelles, called "P granules” in worms, concentrate in specific cells of early embryos and are critical for the normal development of those cells. Cliff Brangwynne et al. later demonstrated that P granules are liquid-like condensates, becoming the first example of a growing list of such condensates that serve diverse roles in eukaryotes.

Finally, Bill himself studied how an organism develops its left–right axis, which in worms becomes apparent when embryos have only six cells. He documented that one-cell embryos already show signs of knowing left from right. From then on, the license plate on his car read WORMS1.

Throughout his career, in part inspired by his educator parents, Bill was passionate about improving science education. He coauthored the textbook Biochemistry: A Problems Approach (1). Later, working with other education luminaries like Bruce Alberts and Jo Handelsman, Bill championed transforming science education away from passive student listening-looking-studying to inquiry-based discussion and hands-on “active learning.” He embraced a classic principle of instruction from Confucius: “I hear and I forget. I see and I remember. I do and I understand.”

Toward the end of his career, Bill turned his full attention to research on science education. He and Jenny Knight tested the effectiveness of incorporating active-learning; students used “clickers” to answer multiple-choice questions designed to gauge their understanding of new concepts and animate discussions with their peers. The data showed that peer discussions and clickers led to significantly improved learning gains. Bill helped spread active learning and the use of clickers from his own classrooms to the international science education community through workshops, his participation in the NAS Summer Institutes for Undergraduate Education in Biology, and his work with the CBE-Life Sciences Education journal. Rachelle M. Spell of Emory University said, “Bill is not only an excellent educator himself, he helped start a revolution across STEM teaching.”

Bill’s life was greatly enriched by his German-born wife, Renate, an accomplished poet and teacher of creative writing. Bill met Renate while traveling abroad during his college years, when she was working as an au pair with a family in Dijon, France. Smitten, Bill began learning German and married Renate two years later. To Bill’s great joy, his passion for music infected his and Renate’s two sons, Oliver and Chris, who have become internationally renowned musicians, performing as The Wood Brothers.

Although Bill left the music career to Joan Baez, he continued to play guitar, compose, and perform songs, often activist ballads, up until his death. He was wildly applauded after his memorable performance of “The Rime of the Ancient Worm Picker” at the 2019 International C. elegans meeting. His innovative songs of social protest and commentary on the human condition delighted his neighbors in the retirement community where he lived for the last six years of his life. He also enjoyed joining his sons on stage when they performed in Boulder.

Bill left a profound mark on science and science education and inspired trainees and colleagues to follow in his footsteps, ensuring that his legacy will live on. His love of music lives on in his sons and the many friends who recall his songs with a smile.

Author contributions

S.S. and J.R.M. wrote the paper.

Competing interests

The authors declare no competing interest.
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1 W. B. Wood, J. H. Wilson, R. M. Benbow, L. E. Hood, Biochemistry: A Problems Approach (W.A. Benjamin, Inc.). ed. 1 (1974), ed.2 (1981).
