School of Aquatic and Fishery Sciences

In 1991, when I walked out of my last final exam in my final year at the University of Victoria, I cold-called my intended PhD supervisor, Dr. Tom Quinn. I gave a long, reasonably well-prepared spiel about my passion for salmon and my desire to do graduate work in his lab. A modest silence followed my monologue and then a “Well, it sounds like you would make an excellent graduate student but, unfortunately, you missed the application deadline by six months.” Momentarily crushed, my enthusiasm recovered when he suggested that I come work for him over the fall. Thus began a seven-year stint with Tom at the School of Fisheries, starting with a fall working on chum salmon at Kennedy Creek in Washington, then a winter working with sockeye salmon fry exiting the Cedar River in Seattle, then a summer in Alaska working with the Fisheries Research Institute (FRI) camps—Wood River (at that time lead by Dr. Don Rogers), Lake Nerka, and Iliamna.

The next year, I met the application deadline, applying at the same time for a graduate scholarship from the Natural Sciences and Engineering Research Council of Canada (NSERC). In the spring, I received a letter from NSERC denying me the option of taking my MS scholarship to Washington University on the grounds that it didn’t have a very good fisheries program. I wrote back politely—but without much hope—to first agree with NSERC that Washington University indeed was not well known for its fisheries program, but that the University of Washington was—and that it was the latter at which I wished to pursue my studies. All was well regardless as I received an H. Mason Keeler scholarship that enabled Tom to take me as a student and, a few months letter, I received a letter from NSERC saying, effectively, “Oops, sorry, our mistake. Here is your scholarship.”

Having had a formative and inspiring set of field experiences in Washington and Alaska, I suggested to Tom that I do my MS on topics similar to those projects on which I had been working. Tom, as always, listened politely and then suggested I instead work on rapid evolution in Lake Washington sockeye salmon. This suggestion turned out to be exceptional as it started my path to being one of the forerunners—along with my office-mate Mike Kinnison (MS, 1997; PhD, 1999) —in the study of rapid evolution. At the same time, I met the great, and ever enthusiastic, Fred Utter who helped me do my first genetic work with allozymes—and still, sadly, my only hands-on genetic work. Of course, all was not always smooth sailing, especially when the boat, the Nettie H, I had worked on for the FRI test fishery in Bristol Bay, sank a few months later while crab fishing, tragically causing the death of all on board.

Having had my MS project suggested to me by my supervisor, I decided I needed to do a PhD all on my own. I therefore suggested to Tom a project at Pick Creek, Alaska, on the reproductive energetics of Pacific salmon. Then followed two extremely intensive summers of field work at the Lake Nerka camp, not only conducting research, but also having a wonderful time experiencing and photographing nature.

The 1995 field season was particularly memorable for probably 50 bear encounters, most of them pleasant and inspiring, but some of them rather scary. I continued to work at Lake Nerka until 2000, even after graduating, making it an even 10 summers of Alaska work with FRI. These years included the first research visits to Lake Nerka by Ray Hilborn and Daniel Schindler, both of whom still work there.

The UW School of Fisheries, now the School of Aquatic and Fishery Sciences, was an outstanding experience for me. I am especially appreciative of my supervisor Tom Quinn, who gave me some great ideas, shaped my manic approach to manuscript editing, encouraged me to explore collaborations with others independent of him, and had a knack for filling his lab with an exceptionally synergistic and energetic group of students. Especially formative for me was having my desk directly beside Mike Kinnison, now a professor at the University of Maine, for seven years. Although we played Doom and Doom II with a serial cable linking our computers between 10 pm and 1 am, we actually did research for at least as many hours before that.

I began graduate school at the College of Fisheries in 1971 after graduating from Penn State. I had worked in the genetics lab of Jim Wright at Penn State, and he recommended that I attend grad school at UW because a former student of his (Bill Hershberger) had recently joined the Fisheries faculty.

The College of Fisheries was not a good place to study genetics in the early 1970s. Don Hagen, then Curator of Fishes, was an outstanding ecological geneticist working on sticklebacks. I remember talking with Don when I was a new graduate student. He felt that his basic science was not appreciated by the College. At the time, Don and George Brown were co-supervising the research of another Fisheries graduate student, Mike Johnson (MS, 1970), who studied the genetics of crested blenny in Puget Sound. Mike later received his PhD at Yale, and he went on to an extremely successful career in evolutionary genetics. Mike has been at the University of Western Australia since 1976.

My early studies were frustrating. I remember phoning Jim Wright at Penn State and telling him that things were not going well. He suggested that I pay a visit to Fred Utter at the nearby Montlake Lab of the National Marine Fisheries Service. After talking with Fred for less than an hour, he offered me a space in his lab and a desk. I literally began working with Fred the next day, and, as they say, I never looked back.

I spent a fantastic and fun four years working with Fred. His support and enthusiasm were tremendous. Allozyme electrophoresis had just been developed, and it made it possible to study the genetics of natural populations for the first time. Those were exciting times! In 1971, Allyn Johnson (PhD, 1972) and I were the only grad students in Fred’s lab. We would later be joined by Bernie May (MS, 1975) and Jim Seeb (MS, 1982; PhD, 1987) before I graduated in 1975.

My difficulties with the College of Fisheries persisted because Fred could not be my primary supervisor because he was an affiliate faculty member. Doug Chapman, the Dean of Fisheries, was extremely helpful to me during those difficult times. Eventually, Fred and I obtained funding for my studies of steelhead from the Washington Department of Game. This funding came through the Washington Cooperative Fisheries Unit under the supervision of Dick Whitney. Dick became a member of my committee, and he was extremely helpful to me during my graduate school career.

I received my MS in Fisheries, but I did not want to receive my PhD in Fisheries because I was primarily interested in genetics. Joe Felsenstein in the Department of Genetics suggested that I apply for an Interdisciplinary Individual PhD under his co-supervision. I was successful and received my PhD in Fisheries and Genetics in 1975.

My down-and-up graduate school career at UW worked out very well for me. I did a post-doc at Aarhus University in Denmark in 1975–76, and I have been at the University of Montana since 1976. Today, there are a number of world-class geneticists on the faculty of SAFS. Back in the 1970s, we were excited at being able to detect a few polymorphic gene loci. Today the geneticists at SAFS are studying whole genomes of many species!

I admit that my undergraduate experience was underwhelming. However, eventually (and fortunately) I found my way to the University of Maine to study for an MS degree in Zoology. In Maine, there were two seminal developments for my career: I discovered shellfish aquaculture and was fortuitously appointed as research assistant on a project to make triploid salmon. In time, these two paths merged, and I was integrally involved in the creation of the first triploid shellfish—oysters, clams, and scallops.

While I was there, the Maine aquaculture industry was being invented. These “downeast” types were conjuring up new ways to grow salmon, oysters, clams, mussels, seaweed, and the like. I was able to experience their growing pains and appreciate the value of practical research on questions of profitability. That appreciation has lasted my whole career.

At that time, the Maine aquaculture industry was not interested in triploids. Creating triploids is a value-added process, and Maine growers were most interested in keeping their oysters alive. I was therefore ready to swear off triploid shellfish. However, one day, standing on the shores of our cottage on the Damariscotta River, I wondered where my future lay, and I realized I needed a research license! I started looking around for a PhD program that combined aquaculture and genetics. I made a grand circuit of potential schools including UW. There were more logical places for me to go, but there was something about UW Fisheries that felt like home. Plus, Ken Chew, the iconic shellfish aquaculture pioneer, was there. UW Fisheries must have had the same effect on other New Englanders. There were four other graduate students also from the Bay State (Massachusetts).

Having sworn off shellfish, I immediately immersed myself in the salmon species of the Pacific Northwest, subjecting several of them to triploid induction. My abstention from shellfish research did not last long. Through fellow grad student Sandra Downing (MS, 1987; PhD, 1993), I learned that there was a large-scale oyster aquaculture industry based on hatchery production in the Pacific Northwest—the exact template needed for integration of a genetic improvement such as triploidy. I wrote a proposal to Washington Sea Grant, and it was funded. What followed for my graduate studies at UW was a brilliant exercise in practical shellfish research, working directly with the oyster industry—Coast Oyster, Taylor Shellfish, Wescott Bay Shellfish—to commercialize polyploid technology. Also overlapping my stay was fellow grad student Ximing Guo (PhD, 1991), doing pioneering research on induction of tetraploidy in oysters.

There was a cadre of grad students in Fisheries who were interested in genetics. We named ourselves the “old gonads,” since several of us were beyond the typical cohort age and took courses in Recombination and Mutation, Molecular Genetics (when the word “molecular” was still novel), Chromosome Behavior (with Larry Sandler), and Population Genetics (with Joe Felsenstein), both renowned in their fields. Better yet, many of us had the benefit of the mentorship of “the founding father of fishery genetics,” Fred Utter, and his extraordinary stable of graduate students and alums, like Fred Allendorf. My first official meeting with Utter was on the back of his 350 Honda motorcycle taking a trip to Goldies for hamburgers, beers, and science.

With this background from UW, I was able to land a faculty position with Rutgers University at the Haskin Shellfish Research Lab in the late 1980s, where Ximing joined me after a few years. At Rutgers, Ximing and I invented tetraploid oysters, which have had a worldwide influence on shellfish aquaculture. After 10 years at Rutgers, I moved to the Virginia Institute of Marine Science as the founder and director of the Aquaculture Genetics and Breeding Technology Center (ABC). ABC is arguably the largest oyster breeding program in the world, and we provide broodstock improvement and management for a large segment of the industry on the East Coast. That magical, formative period during my tenure at UW was, hands down, the most influential in my career.