I am deeply honored by this award, and I feel very privileged to be numbered with its previous recipients.
I want to tell you just a little about my journey as a behavior geneticist and to say a few words of appreciation for my mentors, colleagues and friends. But I will not make an Academy Awards speech thanking everyone from my mom to my cat.
When I went to the University of Chicago in 1956, I was interested in the biological effects of radiation. My interest in this subject may have been due to an overexposure to movies on radiation, mutations, giant insects, and rapacious reptiles. Regardless, I had done a science fair project in high school on the mutagenic effects of X-rays on Dobzhansky's favorite organism, the fruit fly. This was my first experience with sex chromosomes. But my interests changed during the first two years of college. One of my initial courses was about psychology, anthropology, and sociology. We read Freud, Mead, Marx, and many others. I was amazed and excited by their ideas on human behavior. But this did not seem like science to me. No experiments; no numerical data. The following year I took a course in biology. In that class, we read Mendel. Also, we did gene mapping with fruit flies. My next encounter with the sex chromosomes. I had found something that I was good at and that I enjoyed very much. Genetics. Later we read, Darwin's, The Origin of Species. In chapter eight, he argues that behavior evolves. I knew that since behaviors had evolved, genes must have effects on behaviors, and thus, it seemed to me that genetics could be a foundation for a science of behavior. Little did I know that there was at The University of Chicago a professor who had come to a similar conclusion. In the 1950s, he was already teaching a course in behavior genetics.
This professor was Benson Ginsburg, my mentor, colleague, and friend. Our association began in the Fall of 1959. I am greatly indebted to him. Both in where we agree and disagree, he has shaped my views of and approaches to behavior genetics. His pioneering work in the genetics of audiogenic seizures and of aggression were a base for my own research. Over the years, he has been unstinting in intellectual and emotional support. A thank you would barely express my deep appreciation of and regard for Ben Ginsburg.
While we were at Chicago, Joanne Jumonville, a doctoral student of Ginsburg's, observed reciprocal hybrid differences in aggression, and Ginsburg suggested that this was an effect of the Y chromosome. At the University of Connecticut, these findings were replicated and extended by a graduate student, Michael Selmanoff. Selmanoff, Jumonville, Ginsburg, and I published these in a 1975 article in Nature. Soon after, David Hay wrote a letter to Nature suggesting that the reciprocal difference was due to opposite effects of autosomes and maternal environment. This possibility had been modeled previously by David Fulker.
Because of a meeting that I went to as a graduate student, I was able to understand David Fulker's paper and reply to David Hay's suggestion. During the summer of 1961, Jerry Hirsch held a meeting on Behavior-Genetic Analysis at The Center for Advanced Study in the Behavioral Sciences. I was privileged to be allowed to attend that meeting, and I had my first visit to America's spectacular West. At the meeting, I met and heard Jan Bruell, Peter Broadhurst, and Gerry McClearn. They all gave lectures presenting concepts and methods of quantitative genetics as applied to behavior and its evolution. I had never heard of Matherian or genetic triangles, additive variance, diallel crosses etc. But I was very fascinated. One day, I asked Gerry McClearn how I could learn more about this. He told me of a new book. This was Falconer's, Introduction to Quantitative Genetics. When I returned to Chicago, I immediately purchased the book for $6. I read it from cover to cover. It was a great help in understanding the fast growing literature on quantitative genetics and behavior. This included Janet Hyde's selection study of female aggression, Marty Hahn's diallel cross research on male and female aggression, and David Hay's work with triple test crosses and aggression. Also, this introduction to quantitative genetics is now a firm foundation in my search for QTLs with effects on aggression. My great thanks to Jan and Peter but especially to Gerry.
We initiated the breeding of Y congenics in response to David Hay's letter. By 1979, we were able to show with these and some F1s that there was indeed a Y effect in interaction with autosomes on aggression. I consider this paper a classic. This was published in Behavior Genetics. Norm Henderson was then associate editor. At a meeting on behavior genetics in Banff, Canada, he helped me to revise the manuscript. Norm many thanks for this as well as much else.
Much has been learned since then about the Y chromosome and behavior, thanks to my students, Paul Shrenker, Tim Plat, Sonoko Ogawa, Anne Didier-Erickson, and Ed Monahan, and to my colleagues, especially Pierre Roubertoux, Michelle Carlier, Pascalle Guillot, Geert van Oortmerssen, and Frans Sluyter. Another, here tonight, is Kunio Yamazaki who collaborated with us on effects of the Y on urinary odortypes. Also, ideas of Marty Hahn, John Fuller, and Paul Scott on social genetics contributed much to a recently published experiments of Ed Monahn's and mine relating such odortypes to aggression.
In 1986, I took a sabbatical in the laboratory of Mike Dewey at the University of South Carolina to learn the new molecular genetics. Mike and I had collaborated on a study of audiogenic seizures with chimeric mice. Some of you may remember that Mike gave a talk on transgenic mice at our 1989 meeting. I went to Mike's laboratory because he was working on the molecular genetics of the Y, and because Tim Plat was doing a postdoc in his lab. Tim, who had been my graduate student, was now my teacher. We worked on a study of phylogenetic changes in repeated sequences on the Y. A most satisfying moment occurred when the developed film showed that from soups to nuts or DNA extraction to probe hybridization, I had successfully soloed my first Southern blot. This was one of the most enjoyable periods in my career as a scientist. There was also extra frosting on my sabbatical cake; I had the opportunity to indulge in my hobby by visiting all the major and most of the minor Revolutionary and Civil War sites in North and South Carolina.
As a result of my sabbatical, I organized a BGA symposium and edited a special issue of Behavior Genetics on the Y chromosome. There, I argued that genes were added to and lost from the Y during its evolution. This rather than a waning Y is now the widely accepted view of the evolution of the mammalian Y.
Also, it prepared me to collaborate recently with Christoph Pilgrim, Ingrid Reisert, and Gerogia Lahr at the University of Ulm on the expression of Sry in the brain. Sry is transcribed in adult brain of male mice, and it is now a candidate for a gene on the Y with effects on behavior. This is also the basis for my more recent speculation that the Y may have non-hormonal effects on sex differences in brain and behavior and for discussions about this possiblity in humans with Sheri Berenbaum.
I could go on but I won't. There are many , many people who have benefited
my work or touched my life. I thank you all for an exciting, stimulating,
challenging, and rewarding journey in behavior genetics. As I had hoped
and believed in my days as an undergraduate, genetics has become and is
a firm foundation for a science of behavior and mind. Also, Tack (thanks
in Swedish): To Lee Ehrman, Jim Wilson , and Tom Bouchard of the awards
committee. And yes, to my mother, Louise, and my cat, Casey. And especially
to my fiancee, Patricia, who is here with me tonight.