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II. Graduate Fellowship at University of Texas at Austin; Beginning work with Drosophila

AM: OK. Well, why did you decide to switch to biology and chemistry from physics?

JC: I don't know. Not any particularly good reason. I suspect it was the circumstances of enrollment that particular year, and things like that. I know if I'd gone on with physics, I'd be a very happy physicist. I sort of liked the subject.

JC: Well, then I didn't know enough, and the course never did get into the interesting parts of physics. There was no relativity, no quantum mechanics, or anything like that. All I know about that subject I've gotten just by reading later on.

JC: It was [Edmund C.] Sinnott and [Leslie C.] Dunn, [Principles of Genetics] was the standard textbook at that time.^{^[1]} The course, taught by my father of course consisted essentially of reading that book pretty much in order. And I loved it. I was perfectly happy to go further into genetics. I thought of medicine a few times during college and decided not to for two reasons. One is, I don't think I could have done it financially, which essentially made the decision. But by that time, I decided I'd rather go to graduate school anyhow, so when it came time to continue on – I don't think I could have – those were days when it was almost impossible to find a job, and in a sense, graduate school was the course of least resistance. If I got a fellowship, at least I was secure for the next few years.

AM: Was it competitive then? Were other students like you trying to get into graduate school?

JC: Yes. Two or three of my closest friends got into graduate school. Not the same one that I did. So actually, it wasn't too hard, at least for me – only the ones who were pretty good students tried this, of course. This is not very relevant to anything, but I remember it. This college required a course in religion in order to graduate, which I didn't really want to take. I was already slightly rebellious. Quietly, not publicly. But anyhow, I found the course I wanted to take, which was the history of the Christian church. It actually included the Jewish church, too. That history is essentially the history of Europe and I chose the course for that reason, and I'm glad I did. I learned quite a bit of history.

In that class – it was about ten people, I suppose – three of us were science majors and the rest were preachers, or future preachers. The three of us got far and away the best grades. I guess if you've learned the bones of the ankle and the Krebs Cycle,^{^[2]} it's no trouble remembering the names of the Popes. (chuckles) I've been amused by that ever since, that the three of us who were not interested in religion got by the best grades in this course in religion, in church history.

AM: When it came to deciding about going to graduate school, what area did you want to –

JC: I considered both biology and chemistry, and I wasn't quite sure which direction I wanted to go. I have an interesting history that Jim [James D.] Watson^{^[3]} and I share. We don't share very much, but we share one thing. We both were turned down by both Harvard [University] and Cal Tech [California Institute of Technology] for entering graduate school. I applied to a dozen or so schools and, as I said, got turned down by those two. I was accepted to a few places. But the first acceptance was from University of Texas [at Austin], and about that same time I got an offer from [University of] Wisconsin [at Madison] – there were two kinds of fellowships at Wisconsin. There was a good kind and a not so good kind. One paid three hundred dollars and the other five hundred dollars. Well, I got the three hundred dollar scholarship. If I'd have gotten the five hundred dollar scholarship, I think I would have gone to Wisconsin, but I didn't. So I went to Texas instead. Now, if I had gone to Wisconsin, it would have been in biochemistry.

JC: Just because that's where this particular fellowship would have been. And if I went to Texas, it would have been in biology, or genetics. By that time, I think – I wasn't unhappy that I made this choice, and I have never regretted since, either. Also, the Texas offer came first, and I was so insecure that I accepted it almost immediately. I expect if I had been offered at Wisconsin, I'd have accepted that first and would have been a biochemist.

AM: OK. Well, let's talk a little bit then about going to Texas. What were some of the other places you said you got accepted to?

JC: Iowa was one, the University of Iowa. Eleanor Carothers was at the University of Iowa. She knew my father of course very well, and that may have had something to do with them offering me a place there. And I would have studied grasshopper cytology if I had gone there, sort of followed my father's path. Those are the only ones I can remember now.

AM: Now, were you identifying places by their programs in genetics? Texas had just started their program.

JC: Texas was strong in genetics. I'd pretty well decided that if I didn't go into biochemistry – I had almost decided that really genetics is what I wanted to do. I was nominated by my college for a Rhodes Scholarship,^{^[4]} and I went to Lawrence, Kansas, to meet with a group of people. They asked me where I'd go if I really had my choice, and instead of saying Oxford [University], I said Cal Tech [in Pasadena, CA], which was probably the wrong thing to say. But that's how I felt about that time. I felt that's the place I really wanted to go. At that time, the [Thomas H.] Morgan group were all there.^{^[5]}

But Texas seemed good, too, and of course [Hermann J.] Muller was at Texas,^{^[6]} and he'd done his radiation experiments. This would be 1937, and I did know then that Muller had left and gone to Russia, but what I didn't know was that he had no intention of coming back. I thought that while I was there I could probably study with Muller. So I was delighted to get the Texas fellowship. Delighted to get anything, really. I was so insecure about finding employment.

AM: So you were keeping up with the literature in genetics to know that Muller was working on?

JC: I knew that much about it. I was not reading journals. Maybe once in a great while. Most of my knowledge was textbook knowledge at that time. And I had no clear idea at all about the nature of research or that I'd have any real interest in research. I thought I might end up doing pretty much what my father had, go to graduate school, get a degree, and teach.

AM: What kind of laboratory experimentalist opportunities did you have at your college, at the Friends University? Was there much hands-on?

JC: There was quite a bit of hands-on work, but it wasn't research, it was the kind of laboratory you have that goes with courses.

AM: Well, when you got to Texas you mentioned that you wanted to work with Muller. You knew that he was gone, so where were you – Describe the process of who you were going to choose to work with at that point.

JC: Right. Well, when I got there, I wasn't sure who I'd be working with. But the fellowship was not with any particular person. It was a teaching assistantship. But the person who more or less picked me up, I think it would be fair to say, was [John T.] Patterson, and he became my major professor.^{^[7]} He was an interesting character, what in those days you would call earthy. He had a semi-vulgar, funny actually, sense of humor. I remember when I came in, he was gruff. He was always puffing on his pipe and seemed gruff, although he was really kindly. He said, "You're blonder and skinnier than I thought you'd be." Those were his first words.

JC: I had sent a photograph, which you did in those days. He then became a major professor, and I started out just as a graduate student. I was mostly taking courses, and then eventually found my way into the fly lab, but not until a month or so after I'd been there. Mostly I was taking courses, and doing this teaching, which I'd never done before.

The three people who were central to my education were – there was Patterson, as I just mentioned. [Theophilus S.] Painter was the cytologist.^{^[8]} But the one that I interacted most with is a man named Wilson [S.] Stone.^{^[9]}

JC: He'd left by that time. He and Stone were contemporaries, but Stone stayed on as a faculty member. I didn't meet Bentley Glass until several years later.

AM: OK. Now, how was genetics organized at Texas? Did it have its own department at that point, or was it still kind of divided between zoology –

JC: It was part of zoology. There wasn't any genetics department itself. But there was a Drosophila laboratory,^{^[11]} which was all the genetics there was. It was a very popular place, incidentally, because it was a room that was cooled, and in Texas summers, that's the thing you look for, of course. I don't want to say I went into genetics because of temperature control, but that didn't slow me from going that way.

This was an interesting period in the history of genetics at Texas because they were making a transition from studying cytogenetic problems^{^[12]} into studying speciation^{^[13]} and natural populations. I got there just as the change was being made. Patterson and Stone were working on a problem, trying to infer what happened at meiosis^{^[14]} from breeding results. Now it seems like a trivial problem, but at that time, it was difficult for me and for them.

There was also a much more mathematically minded student in the lab. He had taken a crack at this and didn't get anywhere, but I thought I'd look at it. I solved the problem, to my satisfaction. I had one of these – first time I ever had, and it hasn't really happened since. I was stuck on this crazy thing, and I thought about it for two or three days off and on. Finally, I gave up and went to the beer tavern and had a beer and then had a sudden realization of what I had been doing wrong. The kind of thing you read about that happens once in a while. That's happened maybe a couple of other times, but this was the first time and the most striking in my life. So then I went back and solved this problem immediately.

At that time, I hadn't had much math, but Stone hadn't had any math either and was therefore more respectful of it probably than he would have been if he'd had more. He said, "You really ought to take math courses and you ought to read Sewall [G.] Wright,^{^[15]} if you have this kind of mathematical skills." It really wasn't much, but I acted on it anyhow and started – so I then took statistics and probability courses and math courses and started reading Wright, which I found very difficult to read, but I tried at it anyhow. That's all I did in this kind of thing.

Incidentally, I didn't know about maximum likelihood method in those days. That would be the way I would do it now, but I had invented my own little way, which was not nearly so good.

I was more or less assigned what to work on in that Patterson was starting studying different species, and there were three or four different graduate students and each of us got a Drosophila group to work on. Mine was the mulleri group,^{^[16]} so my thesis was on isolating mechanisms principally in that group.

One difference between the attitude that I had, at least, and I think the University of Texas in general – the Ph.D. here nowadays is really a research degree, and you primarily spend a lot of time on research and you're successful or not in accordance with that research. At Texas at that time, there was more emphasis on learning, and the thesis was something you did all right, but my thesis that I wrote then certainly wouldn't pass muster here now. It was done quickly and didn't have very much data. I spent a lot of time taking courses. In a way, I'm glad I did.

As I said, Stone was more influential with me than Patterson was, and he encouraged me – and Patterson didn't disagree – encouraged me to take courses that I wasn't really prepared for. He said, "If you get a low mark, don't worry about it." So I have some B's on my record that one shouldn't have as a graduate student. I took a course in radiation physics, for example, that I was totally unprepared for, because other students were physics majors. But I learned a hell of a lot in this course, and I've been glad I did it ever since. I took physical chemistry, too, and that I thoroughly enjoyed. I might have become a physical chemist if I'd had taken it earlier in life. (chuckles)

JC: Well, I took to it. I even had occasion to use this later. I'll again jump ahead. The book we used was by *[Frederick H.] Getman and [Farrington] Daniels [Outlines of Physical Chemistry].^{^[17]} I remember the authors very well. Daniels was at the University of Wisconsin, and I later met him. It was a pleasure to meet the man whose textbook I'd used. But since I did like it – I worked all the problems in the book. Some year later, after coming to be on the [Wisconsin] faculty, Joshua Lederberg^{^[18]} had a student, Norton Zinder^{^[19]} – he's become very distinguished since that time. He was taking physical chemistry, which he didn't like. He was in your category. (chuckles) But Josh had more or less insisted on his taking it. Then I found out he was using the same book I had, so when he had any trouble with problems, I would help him with those problems. I don't know how much responsibility to take for his getting through physical chemistry and ultimately becoming a big success, but I'll take any amount of credit that I can get. (laughs)

AM: Where there any corn geneticists or any geneticists in the botany department at Texas at this time?

JC: There weren't at that time. There were some corn genetics going on, but it wasn't at Austin, it was at Texas A & M [University]. I don't believe there was any other genetics in Texas then. There was cytology, and there was a course in fungi, which I took, but I think this was all of the genetics. It's an interesting question which I never thought of before. But I certainly didn't get any genetics anywhere else.

AM: OK. How much thought did you give to model organisms, or was that even an issue?

JC: No thought at all. They were using Drosophila and that was it. I enjoyed actually working with Drosophila, so this wasn't any hardship. I've never been the kind of person who goes to the laboratory and just enjoys every minute of laboratory manipulation. I'm really more interested in getting an answer of some sort and thinking about it than I am of the day-to-day details. And that was true then, too. I probably did less work per thesis than any of my contemporaries did.

JC: I wasn't particularly bad, or good either. There wasn't anything that called for laboratory skills. Well, I did learn how to blow glass, for example. After I moved to my first job at Dartmouth, I was the only one in the department who knew how to do this, so I made equipment for other people, but I never thought of it as being a particular skill.

JC: Mm-hmm. I remember my father told me once that, "Whatever else you go into, don't go into Drosophila genetics because you don't want to spend all your time counting millions of flies." Then of course, that's what I did. But I usually managed to find a way of working on a problem that didn't demand an awful lot of counting work.

Let me mention one more thing while I was going to graduate school, because I did keep up music during those graduate school years, which was hard to do because graduate school is a full-time job. But I sort of disciplined myself to keep up by playing in the university orchestra. Also, you could rent a practice room – not rent it, I didn't have to pay for it, but get assigned to a practice room. I did that from five to six every day, for several years. Not all four years, but much of the time. That just meant that I had to go to that practice room every afternoon or else I couldn't have kept it. So that kept me from going totally stale.

I did a little bit of freelance playing while I was there. During my last year before getting a degree, I changed from the teaching appointment to a research appointment, which paid a little bit less. There was an opportunity as a musician for playing in a little orchestra that made up the difference, so I did that. I enjoyed doing it. It was corny and terrible, but I made a little money that way. On the lighter side, I was a victim of one of the diabolically cleverest jokes that anybody's ever perpetrated. I was playing in the pit orchestra for kind of a light comedy show of some sort. I can't remember what it was now. I stopped to get a hamburger for supper and left my viola in the fly lab and came back and started playing, and fruit flies kept buzzing up out of the f-holes. And one of my dear colleagues had anesthetized thousands of flies and put them inside the viola. (laughs)

JC: There probably was. I did play in a dance band for a little while, but I had no contact with the blues, any of that kind of music. It was mostly classical.

AM: OK. Well, you just mentioned that your dad said whatever you do in genetics, don't end up spending all your time counting flies.

AM: And that's what you do. But maybe you can talk about at Texas what techniques and methodologies did you have available to you?

JC: The main technique was a microscope, and I did use salivary gland chromosomes.^{^[20]} I learned to [analyze] them. Other than that, the technique was simply that of growing flies. Sometimes I collected my own flies, but other people were doing collecting and they usually -- so I was interested in studying flies collected from nature. They did the collecting for me essentially, so I got more flies than I could work with.

I did enjoy this. I found four different closely related species that hybridized to some extent,^{^[21]} and I was able to, in some cases, do a genetic analysis of what caused the hybrid sterility. And I found an interesting mutant that had just a slight effect within the species but had a gross effect between species. Nowadays, that's commonplace, but that was a little bit novel at the time.

I also discovered something that's become very much talked about recently. I didn't establish it very well, nor did I realize the significance. It's what is now called reinforcement. That is, if you take two species that are partially incompatible in the same environment, they'll become totally incompatible; otherwise, the system breaks down. And that building up of incompatibility is called reinforcement.^{^[22]} I found that flies from the same locality, different species from the same locality, were very reluctant to mate. Whereas, those from different localities would mate, at least in the laboratory. But I didn't make much of it. My thesis was finally published at the University of Texas publication. They had a series of papers on [Drosophila genetics].

JC: No. It was a year or so later. I wrote it while I was there. I actually did have one paper published while I was still there. I wasn't that excited about doing publication, and I wasn't even sure that I wanted a research career. I liked it, but I wasn't sure that I would be any good at it.

AM: I've interviewed quite a few young scientists, within the first five years of their first faculty tenure track appointment, biomedical researchers, and to them, it would be anathema to think that they could leave the graduate program without having not just publications, but some significant publications, as perhaps judged by where they were published. How different was that –

JC: This is just a different time. When it came time to begin looking – Well, I had hoped to do a postdoctoral fellowship, and I really did want to study with Sewall Wright. Patterson did ask Wright at the meeting if it's possible to work out something, but it didn't work out. I don't think it would have been practical anyhow, because I graduated in 1941 and it was pretty apparent we were going to be in a war before the year was out, and I don't think a fellowship was the right thing at that time. So I wasn't really disappointed not to. I had not even met Wright at that time. I admired him greatly by just what he wrote. I also read [Ronald A.] Fisher, too.^{^[23]}

Then I just decided to look for a job. I applied to quite a number of different places, and the one I got was at Dartmouth College. I believe that was almost the only significant opening that year. There weren't very many graduates, either, but jobs were scarce then. My rival was a good friend then, and still, for the position at Dartmouth. It's interesting why I got that job. It was to replace Jim [James V.] Neel.^{^[24]} Sorry you didn't interview him. He would have been a good one. You're a little too late. It was to replace him while he went on leave for a year. I was reasonably sure he wasn't coming back, but anyhow, the appointment was for one year and nobody knew what would happen after that year.

Another person that was considered for this appointment was Harrison [D.] Stalker, who maybe you have heard of.^{^[25]} His mentor was Jim Neel. It wasn't his mentor, I'm sorry. Neel knew him, though, and recommended him. When I got there, I discovered what the story was. The man that taught elementary biology, which Neel had assisted and then I was to assist, just didn't get along with Jim Neel at all. Well, the long and the short of it is, if Neel recommended someone, he wasn't going to have him. (chuckles) So the reason I got this job was because Neel didn't recommend me. I think the department thought that they didn't have much choice between these two people, and they might as well appease the fellow. So that's why I came.

AM: Before we move on to Dartmouth, I want to talk a little bit more about Texas, and right off the bat, how common was it for graduate students to do postdocs then?

JC: Hardly ever. It certainly wasn't the routine thing that it is now. Maybe I graduated at the wrong time, but I think the people that were ahead of me – the ones that had postdocs, at Texas at least, were the ones that stayed on at Texas because they couldn't find a job. So somebody scrounged around and found an appointment for them. But this was not a postdoc in the sense that you would seek it out, it was a temporizing affair.

JC: Would be really to learn something from him. I would have profited by it, I'm sure, but of course I got him as a colleague later on, which is even better.

AM: You had mentioned that [Wilson] Stone had encouraged you to take a more mathematical statistical approach to a problem as a means to solving the problem. At that time, how important did you think mathematic statistics was going to be in your work?

JC: Well, I just liked it, and so I went in for that reason. I wasn't sure that I would ever be good enough to make significant accomplishments, because it was all I could do to understand either Fisher or Wright, and I had no feeling that I could carry on or do anything [comparable to what] they had done. Mostly to me it was a matter of just enjoying learning it without giving too much thought as to what I would do with it. That sort of came later.

AM: How important was it to read Wright and Fisher among the other graduate students?

JC: I was the only one who did it. I talked with Stone about it, but I was the only one who had any real understanding of Wright's work. Stone and I talked a great deal in general about evolutionary problems, but the truly mathematical part, he didn't understand what I was doing. And I barely understood what was Wright was doing. I kept on even after I moved to Dartmouth reading Wright's papers and spent many evenings doing that. But I'll talk about Dartmouth later.

I can't see much else to say about graduate student life. The group of students who were working in genetics were all good friends, and we rented a house and lived together one year. Each of us had our own thesis we were working on. We were doing pretty much the same thing. So it was a pleasant kind of companionship.

AM: One last question and we'll leave Texas and we'll leave off for the day. That is, the program at Texas when you arrived there, *[Hermann J.] Muller was gone, but they had kind of brought him in to build the department up. Muller is an interesting character for a variety of reasons, and he left to go to Russia, and his political consciousness is part of the kind of mystique of him as a geneticist as well. When you got there, what was the legacy at that point being formed around Muller's tenure at Texas?

JC: Well, he certainly left a legacy, and later on I can tell you quite a bit about him because I got well acquainted with Muller later on. But what did I know then? I knew that he had been leftist, maybe Communist, I didn't know for sure. And I knew that he'd gone to Russia for idealistic reasons. And I knew that he had been associated with various kinds of leftist student politics at the university and had essentially burned his bridges. I discovered it after getting there, I didn't know it till I got there. But then I soon learned he wasn't coming back.

There was a lot of talk about Muller. He was certainly the most famous of the Texas people. It was clear enough that Patterson didn't like Muller, but he respected him. And Stone had been Muller's student. He both liked Muller and respected him. But neither Patterson nor [Theophilus S.] Painter really liked Muller. Muller thought they were stealing his ideas, and I think that's – stealing is the wrong word. They were using them [freely] without [thinking about] stealing.

The problem that Patterson was working on, just finishing up when I got there, was one that was suggested by Muller. I'll tell you what it was. There was a question, raised by Muller, I'm sure, and that is whether the X chromosome determines femaleness by a single gene on the X chromosome, or whether it's a whole series of genes along the chromosome. Well, Patterson had started working on that problem, I'm sure instigated by Muller, by using translocations^{^[26]} and combining left and right halves of translocations and systematically covering the X chromosome this way.

But there was one region he couldn't get. The flies all died. I remember – and it was a suggestion from Stone, the technique for this – that instead of using translocations, to start out with a fly that had a long duplication, which included that region and wouldn't have lived with such a long duplication, but to radiate the fly, and hope to delete much of this [chromosome] and leave just the right region. So I worked on that, and it worked. And I finally solved this problem; when the fly came through, it was not a female. No, it was a male. Which means there was not a single female sex determining system. I did that while I was still a graduate student. I didn't write it until after I got out, but it was one of the first papers that I wrote from Dartmouth. That finished up this problem, but to answer the question you raised initially, this was the Muller heritage, and clearly this was a problem that had been suggested by Muller.

I know from later conversations with Muller, but even more from reading, that he felt that the group in Texas was simply capitalizing on his ideas, which was true. But I've never thought that was so bad. Muller had far more ideas than he could carry through himself. But Patterson clearly was – although, as I said, he really didn't like Muller, but he respected him.

Painter was working on Muller's ideas because, with the salivary gland chromosomes, clearly the idea of using them was a way of discovering the fine structure of genetic mapping. ^{^[27]}That came after Muller had left. Well, I can't be too specific about what Muller's interests on Painter were. Even before he worked on salivary gland chromosomes, he did the same kind of question with other chromosomes, essentially to show that when chromosomes broke, the genetic consequences were what you would expect from that.

AM: Did his persona as activist-scientist [transmit to] any of the graduate students?

JC: Well, I would have been receptive. But there was no political activism among graduate students, among us graduate students at least, at Texas. I tried a little bit to join some societies, but nothing came of it. So essentially, I was just a graduate student and forgot all my political activism. There were stories about Muller and about his leftist views. Patterson was right wing in his own views, so he didn't approve of Muller at all in his politics. Stone, I think, was sympathetic, and he'd been a student of Muller.

They had left by the time I got there, but Muller had two Russian people come to work with him. One was named [Israel I.] Agol and the other was named [Solomon] Levit.^{^[28]} They had left at the same time, or before Muller left actually. I remember several people talking about these Russians there who kept trying to propagandize and proselytize these people and convert them into Communism. But that's the wrong thing to do in Texas (chuckles), and I don't think it got anywhere. The students I was with came later, so they didn't know any of this, but among the faculty members and the people who had been around a long time, most of the talk about these two Russians was the gossipy and amusement sort, rather than whether they were influenced by the politics.

One of the things that distressed me is that both of those two were liquidated pretty shortly after they went back. They were both charged, before they were killed, by being anti-Communist. Well, they certainly weren't when they were in Texas. They made a nuisance of themselves at Texas by preaching Communism.

[1] Edmund C. Sinnott and Leslie C. Dunn, along with Theodosius Dobzhansky, published Principles of Genetics: An Elementary Text with problems, in 1925. The book went through three subsequent editions.

[2] Krebs Cycle: An intricate physiological process with multiple steps through which organisms use oxygen to produce energy and CO₂. Also known as the Citric Acid Cycle and the Szent-Gyorgyi-Krebs cycle, it was elucidated in the 1930s by Nobel laureates Albert Szent-Gyorgyi (1893-1986) and Hans Adolf Krebs (1900-1981).

[3] James D. Watson (1928 -) and Francis Crick (1916-2004) were the first to propose the double helix structure of DNA in 1953 and are renowned for having done so. Along with Maurice Wilkins, Watson and Crick received the 1962 Nobel Prize for their work. See http://www.time.com/time/time100/scientist/profile/watsoncrick.html

[4] Rhodes Scholarship: A prestigious scholarship awarded annually for international students to study at the University of Oxford.

[5] Thomas H. Morgan, (1866 – 1945), A leading American geneticist awarded the Nobel Prize in Physiology or Medicine for his work explaining the locations of genes on chromosomes and the mechanism by which they transmit hereditary traits in his work with Drosophila at Columbia University. See http://nobelprize.org/nobel_prizes/medicine/laureates/1933/morgan-bio.html. Morgan’s group included his students Alfred H. Sturtevant and Calvin Bridges, who arguably contributed at least as much to the work as Morgan himself. With Hermann Muller, the three authored The Mechanism of Mendelian Heredity in 1915.

[6] Hermann J. Muller: (1890 – 1967), A leading American geneticist and former student of Morgan’s, who was awarded the Nobel Prize in Physiology or Medicine in 1946 for demonstrating that artificial mutations can occur as a result of X-ray manipulation. See http://nobelprize.org/nobel_prizes/medicine/laureates/1946/muller-bio.html.

[7] John T. Patterson, (1878-1960), distinguished American researcher in Drosophila (fruit fly) genetics and embryology. He worked with Hermann Muller to show that mutations were reversible.

[8] Theophilus S. Painter, (1889 – 1969) American zoologist who worked with Muller to identify Drosophila genes and was the first to come close to a correct approximation of the number of human chromosomes.

[9] Wilson S. Stone, (1907 – 1968), American geneticist who worked with Patterson, Muller, and Painter to examine the effects of ultraviolet radiation on mutation.

[10] Hiram Bentley Glass, (1906 – 2005), leading American biologist and pioneering geneticist who was noted for his writing on the dangers of eugenics and the nature of science in the future. Served as President of the American Society for Human Genetics in 1967. See http://www.nytimes.com/2005/01/20/science/20glass.html. His papers are at the American Philosophical Society; See:

http://www.amphilsoc.org/mole/view?docId=ead/Mss.Ms.Coll.105-ead.xml;query=Bentley Glass .

[11] Drosophila: A genus of fruit fly, most commonly Drosophila melanogaster, that has become a model organism used in genetics experiments since it is relatively inexpensive, easy to care for, has a high fecundity and a short life span, and exhibits a readily discernable phenotype and genotype. See the NCBI site at: http://www.ncbi.nlm.nih.gov/mapview/map_search.cgi?taxid=7227 .

[12] Cytogenetics: The genetic subfield that focuses on the structure and functions of the cell and its chromosomes.

[13] Speciation: The evolutionary process by which new biological species develop.

[14] Meiosis: Cell division producing gametes or spores, leading to sexual reproduction.

[15] Sewall G. Wright, (1889 – 1988), leading American geneticist who helped to develop theoretical population genetics. He computed the amount of inbreeding of members of populations as a result of random genetic drift and, with R. A. Fisher, pioneered methods for computing the distribution of gene frequencies among populations. His papers are at the American Philosophical Society: See http://www.amphilsoc.org/mole/view?docId=ead/Mss.Ms.Coll.60-ead.xml.

[16] Mulleri Group: A subspecies of Drosophila.

[17] This noted textbook has gone through multiple editions under three names and at least four different authors . The first was published by Getman alone in 1913 as Outlines of Theoretical Chemistry; Daniels joined him on the 5^th edition (Outlines of Physical Chemistry) in 1931 and continued publication after Getman’s death in 1941. Crow is presumably referring to the 6^th edition (1937).

[18] Joshua Lederberg, (1925 – 2008), American molecular biologist, who demonstrated that bacteria can exchange and transmit genetic information in reproduction. He received the Nobel Prize in Physiology or Medicine in 1958. One of the deans of American science, he served on numerous committees and also worked with artificial intelligence and space biology programs. See http://www.nobelprize.org/nobel_prizes/medicine/laureates/1958/lederberg-bio.html .

[19] Norton D. Zinder: (b. 1928), American biologist who discovered in 1952 the process of genetic transduction, the use of bacteriophage (a virus) to exchange genetic information between organisms in Salmonella bacteria.

[20]Salivary Glands: ducts in mammals that secrete salivary amylase to break down food in the mouth. Geneticists study salivary gland chromosomes in Drosophila because they are relatively large and easy to see.

[21] Hybridized, that is, cross-bred across species or subspecies.

[22] Reinforcement: also known as the Wallace effect, after evolutionary pioneer Alfred Russel Wallace (1823-1913), is the process by which natural selection increases reproductive isolation through the increased incompatibility of species and subspecies until they can no longer cross-breed or produce only sterile or unfit hybrids.

[23] Ronald Aylmer Fisher, (1890 – 1962), great English statistician and geneticist who created many of the tools and theories of modern biostatistics.

[24] James V. Neel, (1915 – 2000), American geneticist who was one of the first to define genetics as its own field of study. He researched genetic epidemiology, the genetics of sickle cell anemia and the effects of radiation from atomic bombings. In 1956, Neel was the founder of the first US Department of Human Genetics based in a medical school at the University of Michigan. See http://www.aps-pub.com/proceedings/1461/109.pdf.

[25] Harrison D. Stalker, (1915-1982), American evolutionary biologist and late professor at Washington University in St. Louis.

[26] Translocations: the movement or rearrangement of sections of chromosomes, leading to gene fusions and other types of abnormality. Translocations can be detected through cytogenetic studies (karyotypes).

[27] Genetic mapping: Refers to a type of gene mapping using linkage analysis to determine the relative positions of different genes on a chromosome, based on the frequency of their recombination.

[28] These Russian geneticists and students of Muller’s were executed by the Soviet government in the late 1930s.

[1] Neurospora: a genus of fungi. The bread mold species Neurospora crassa is often used as a model organism in genetics; it reproduces rapidly and has minimal nutritional needs. Specifically, Neurospora was used by George Wells Beadle and Edward Lawrie Tatum to develop mutations differing in specific nutritional requirements; this work led to their Nobel-winning (1958) one gene-one enzyme hypothesis.

[2] Archibald E. Garrod, (1857 – 1936), British physician who was the first to claim that genes . He worked extensively on hereditary metabolic disorders, for which he developed the term, “inborn errors of metabolism,” and postulated that these resulted from genetic mutations resulting in the deficit of a single enzyme. His landmark work, Inborn Errors of Metabolism, was first published in 1909.

[3] Endocrinology: The study of the endocrine system, the system of glands and tissues secreting endogenous hormones that regulate the body’s growth and metabolism.

[4] Dr. Royal Alexander Brink, (1897-1984), plant geneticist who was the head of the Department of Genetics at the University of Wisconsin-Madison from 1939 to 1951.

[5] DDT (dichlorodiphenyltrichloroethane) is a controversial pesticide first synthesized in 1874. It was used very effectively to control malaria and typhus in World War II and subsequently much used in agriculture and as a disease prevention measure in the US and in the 1950s. The 1962 book, Silent Spring, by the biologist Rachel Carson, brought attention to its lethal effects on bird populations. DDT’s routine use in agriculture was banned in the US in 1972, and worldwide by the Stockholm Convention of 2001, but it is still in limited local use for disease control.

[6] Second-generation Japanese-American.

[7] Now Professor Emeritus of Zoology at the University of Oklahoma.

[8] Newton Morton is currently Senior Professorial Fellow in Human Genetics at the Human Genetics Division of the of the University of Southampton School of Medicine. His major contributions have been in the fields of human genetic linkage analysis and genetic epidemiology.

[9] Motoo Kimura, (1924 – 1994), A very prominent theoretical population and evolutionary geneticist who proposed in 1968 that most genetic change at the molecular level is neutral with respect to natural selection – that is, genetic drift is a primary evolutionary factor. Crow has written memoirs of appreciation for Kimura in different scientific journals. For one such article, see Crow, JF, “Memories of Motoo.” Theor Popul Biol 49 (1996): 122-127.

[10] Logarithm of the Odds Score, a measurement of the probability that a set of genes are linked, developed by Newton Morton.

[11] Jay Lawrence Lush, (1896 – 1982), American animal geneticist based at Iowa State University who advocated selective livestock breeding based on genetic and quantitative information rather than physical appearance and subjective perceptions. Lush is sometimes known as the father of modern animal breeding.

[12] Consanguinity: Kinship based on sharing common ancestors.

[13] Biological Effects of Atomic Radiation Committees (BEAR) were established by the US National Academy of Science in 1954 to compile and make sense of available knowledge and research about the effects of atomic radiation on living organisms. There were six BEAR subcommittees, each studying a particular area of life, and a number of subcommittees, including one on Genetics (1962-64). The committees published a series of reports before BEAR was phased out in 1964.

[14] Morton NE, Crow JF, and Muller HJ. “An estimate of the mutational damage in man from data on consanguineous marriages.” Proceedings of the National Academy of Sciences of the U.S.A. 1956; 42: 855-863.

[15] Rayla Greenberg Temin: American population and Drosophila geneticist perhaps best known for this 1960 paper with Crow on recurrent mutation and inbreeding. See Greenberg R and Crow JR, “A Comparison of the Effects of Lethal and Detrimental Chromosomes from Drosophila Populations.” Genetics 1960; 45: 1153-1168. She was married to Howard M. Temin (1934-1994), co-recipient of the 1975 Nobel Prize in Physiology and Medicine.

[16] John Z. Bowers, (1913 – 1993), American physician who served on the 1949 commission to study the long-term biological effects of atomic radiation in Hiroshima and Nagasaki and pursued research in this field throughout his long career. He served as Dean of the University of Wisconsin School of Medicine 1955-64.

[17] Klaus Patau, (1908 – 1975), German-American geneticist who worked at the University of Wisconsin Madison from the 1950s, and in 1960, found the extra chromosome in trisomy 13 responsible Patau syndrome, a developmental disorder involving multiple defects, including heart and kidney abnormalities.

[18] Oliver Smithies, (b. 1925), British-American scientist who invented gel electrophoresis in 1955 and the same year developed the technique of homologous DNA recombination to alter animal genomes, making possible the creation of “knockout mice” and targeted gene mutations. For the latter accomplishment, Smithies shared the Nobel Prize in Physiology and Medicine in 2007.

[19] John M, Opitz, (b. 1935), German-American medical geneticist and Professor of Pediatrics and Human Genetics at the University of Utah. Opitz was one of the first to identify specific groups of pediatric anomalies as genetic syndromes, several of which now bear his name. He was the founder (1976) and long-time Editor-in Chief of the American Journal of Medical Genetics, and a founding member of the American Board of Medical Genetics.

[20] Curt Stern: (1902 – 1981), pioneering German-American geneticist who taught at the University of Rochester 1933-47 and at UC Berkeley from 1947 until he retired in 1970. He was the first to demonstrate the crossover of homologous chromosomes in Drosophila, and made many contributions in the areas of genetic recombination, radiation risk, gene regulation, and the separation of human genetics from race-based eugenics. His landmark textbook, The Principles of Human Genetics, was first published in 1949.

[21] Eugenics refers to the management of breeding practices, specifically of human beings, to improve the genome. Highly popular and advocated by many distinguished scientists in the early 20^th century, eugenics became associated with forced sterilization, racial discrimination, euthanasia, and Nazi extermination programs in the 1930s and 1940s and geneticists made a conscious effort to distance themselves. Recent developments in prenatal diagnosis and in genetic engineering technology have brought the ethical and social questions involved in genetics to the fore again.

CROW, JAMES

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JUNE 1, 2005 - JUNE 3, 2005

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