From fruit flies to fallout: Ed Lewis and his science

Edward B. (“Ed”) Lewis was born in Wilkes-Barre, Pennsylvania, on May 20, 1918, and educated at the E.L. Meyers High School in Wilkes-Barre. He grew up during the Great Depression; his father, a jeweler and watchmaker, supported Ed's educational and musical aspirations, despite the economic hardships. One of the highlights for Ed was the public library in Wilkes-Barre, which he used to visit daily. Two books and a journal, which he read there as a teenager, had a major impact on his life. The journal was Science and the first book was Bertrand Russell's The Scientific Outlook (Russell,1931), both of which I will return to later. The second book, The Biological Basis of Human Nature by H. S. Jennings, a Professor at Johns Hopkins University, and dedicated by him to Calvin Bridges (Jennings,1930), is where Ed first read about Drosophila and the power of Drosophila genetics (Lewis,2004). Shown in Figure 1 are two illustrations from the book. On the left is “sex-linked inheritance” of the white gene, which was the subject of T.H. Morgan's first paper on Drosophila genetics in 1910; on the right is the inheritance of the dominant Bar mutation, which A.H. Sturtevant did a lot of work on, and which was important in motivating Ed to his graduate work. Ed does not remember seeing the Bar figure and says that, if he did, he probably would not have understood it anyway! Drosophila genetics as illustrated in H.S. Jennings' 1930 book The Biological Basis of Human Nature, which Ed Lewis read while in high school. On the left are the parents and progeny from a cross of white females to wild-type males. Because of sex-linkage, the male progeny are white eyed whereas the female progeny have red eyes. On the right is a cross of males carrying the sex-linked dominant Bar mutation to wild-type females, which results in females with Bar eyes and males with wild-type eyes. Reprinted from Jennings (1930). Ed was a member of the E.L. Meyers High School biology club, which at that time was chaired by Ed Novitski, who also become a Drosophila geneticist. In late 1934, Ed (Lewis) noticed an advertisement in Science for Drosophila cultures from Turtox Products in Chicago for a dollar each. I searched through 4 years worth of Science to find this advertisement but failed. Both Eds, however, assure me that it existed. After a Web search, I did manage to obtain the 1933 issue of a Turtox catalog (Turtox,1933) and sure enough, there the ad is, offering live Drosophila mutants for two dollars a culture (Fig. 2). Ed thinks that, because of the Great Depression, by 1934 the price had dropped to only 1 dollar. Novitski tells me that not being Drosophila geneticists, when he wrote the letter to obtain this culture, he asked whether for the same price they could throw all the mutants into the same vial. Professor Rifenberg at Purdue who filled the order, wrote back saying that it was clear that they were new to genetics, but for the price, he would send several vials. That was the beginning of Ed's career as a geneticist. A 1933 Turtox catalog with an ad for Drosophila stocks. The ad is very similar to the one that Ed saw as a high school student in the Osterhout Library in Wilkes-Barre, Pennsylvania, in late 1934. Lewis and Ed Novitski obtained the stocks for 1 dollar, launching both of them on the road to becoming Drosophila geneticists. [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com]. In his autobiography, Ed emphasizes that “by allowing Novitski and me freedom to use the biology lab and its supplies to carry out our experiments on Drosophila, our biology teacher, who also was the athletic coach, could not have been more helpful in furthering our careers. There was none of the present attitude that one cannot become a scientist without having had the benefit of teachers skilled in the art of keeping their students constantly motivated” (Lewis,2004). In other words, Ed was motivated from the beginning and has stayed motivated throughout his life. Ed is an excellent musician (Fig. 3). He was given a wooden flute by his great-uncle, Thomas Wyllie, in 1928. Later, during the Depression, his father somehow managed to come up with funds to buy Ed a silver flute. He played in the High School Orchestra and the Wilkes-Barre Symphony, beginning his tertiary education at Bucknell College on a music scholarship. Ed playing the flute in the living room of the author's home in Toronto in 1996. He began his tertiary education on a music scholarship to Bucknell College in 1936 but switched to the University of Minnesota after 1 year to focus on biostatistics and genetics. He remained an enthusiastic and accomplished flautist for the rest of his life. Reprinted from Lipshitz (2004). [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com]. After a year at Bucknell, science triumphed and Ed went to the University of Minnesota to complete his undergraduate studies. He chose that university for two reasons: first, money was tight, and the University of Minnesota had the lowest out-of-state tuition of any major state university; second, because the Reserve Officer Training Corps (ROTC) was not compulsory. Ed's older brother, Jimmy, who went on to a distinguished career in the US diplomatic service, assisted financially, enabling Ed to complete his undergraduate degree in biostatistics, which he did in 2 years. It was at the University of Minnesota that Ed started working on the recessive allele of Star, which Novitski had found and had sent to him (Lewis,1939). Ed continued these analyses as a graduate student at the California Institute of Technology (Caltech) as one of A.H. Sturtevant's students. During his 3-year graduate career (1939–1942), Ed was to invent the “cis–trans test” for position effects that all of us study in our introductory genetics courses (Lewis,1941,1942,1945). The Second World War interrupted Ed's career as a geneticist. He stayed on at Caltech to complete a Master's degree in meteorology and then joined the US Army Air Corps, first in Hawaii and subsequently as a weather officer for the US 10th army in Okinawa before the invasion of Japan (Fig. 4). Lieutenant Lewis at his weather officer's desk, G2 section, US Tenth Army, Okinawa, 1945. Reprinted from Lipshitz (2004). Ed returned to Caltech in 1946. In his autobiography he writes that, in 1943, “Millikan called me into his office and simply said: ‘This war will not last forever. When it is over we would like you to come back as an instructor’…Obviously Sturtevant had a hand in this style of making appointments that was typical of the times” (Lewis,2004). Ed has spent his entire independent career at Caltech, becoming the Thomas Hunt Morgan Professor of Biology in 1966, attaining Emeritus status in 1988, but remaining active in his lab. The other big event of 1946 was that Pamela Harrah became Pamela Lewis. Beadle had returned to Caltech from Stanford in 1946 to chair the Biology Division. That same year Ed, as an Instructor at Caltech, had taken over the Drosophila stock collection and was looking for a stock keeper. While still at Stanford, Beadle called Pam (who had done an undergraduate degree there) into his office and said, “Hey Pam, how tall are you?” to which Pam replied, “5' 3”.” Beadle then said, “Your new Boss is 5' 4” tall, he's 28 and maybe you will like him so much, you will fall in love and decide to there at and A after Ed and Pam were and have remained so for over years (Fig. Ed, Pam, and Lewis in the high to the on the in Ed had been a which he chose to in at Reprinted from Lipshitz (2004). Ed's to science is to his as as his Ed chose to from Bertrand Russell's book The Scientific Outlook he had first as a high school his power of is the of and with in the of science are at the of Ed's is an Ed was at home in genetics, and his first love has been genetics. The two in genetics are of the of inheritance and Sturtevant's as an undergraduate at University in while working in Morgan's of of on all of Ed's on Drosophila present his in of to Russell's I not to the but I it be to at a of taken from Ed's over a (Fig. It be that Ed not only but he them on was in other as and of Ed's A for the of by Second from A for with the of the University of Chicago from from to the for in with of Ed's for that a which he to the from the of the It is very important when Ed's to he and he is Ed's of a that he has The in are his of one the which from Ed's of the (Fig. the and are in there is a very when they are in there is a The to this the of the time (Fig. On the left is the which from the results of the for two On the right is Ed's of the In this and would the of the wild-type and while and would the of the wild-type and On the these are in and on the right in Lewis me as I was the to Ed's was years after of these were have very of their a lot about the of but we not also have of the of the for which Ed is that the of the in the is the same as the order, the of the they (Fig. The The of the on the with the of the they Lewis [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com]. Ed's work can be into two first his Drosophila second, his work on and I will first his Drosophila the first years of Ed's career in Drosophila, his were the gene, how it how it and how it Ed's to which is he is that when you you up where you you Ed began this work several years before was to be the and before the of was the were as on like on a It was that only Sturtevant's had that, if two recessive a and a when in they are of the same would not be to a and In in had a at the Bar (Fig. when the two Bar are present in on the same the which is a in the of the is more when the two are present in in the Drosophila in had that was that a which subsequently and their The for Ed, an was to whether it be to obtain of a In how would one Ed's of the for position effects accomplished this He the in his to the and (Fig. to and white and and the beginning in 1946 for of these experiments that in Ed and that the and as as the of to (Fig. as Bridges had be the for The position the and which Ed during his On the left is the which results in an wild-type On the right is the which the Lewis to In and to and the to Lewis Ed to out very the that by two the of the of that the in one of the of a into a new He on to the and on the in The and was years by In the Ed position that he called When he a that would he found that the of two was from that when the were to A is with and (Fig. in this the when is Ed that there be that is to from one to the the In the of and of of this making the more has been we not the of years after Ed the When are to only a of the second can be When the are to a more of the Ed's is the of Reprinted with the of the University of Chicago from Lewis major from Ed's of the the of mutants (Fig. with his this Ed, in his to the that the mutants the of and that the second in is the The of the is to from this state to more from second to as as but of the second by in Ed to the a wild-type in of the of the is to the In in of the of the Lewis The second a The of the is to the more from a second of After Lewis In the Ed a very of that he called and The mutation of the The mutation, in the (Fig. the is which is similar but not to of in Ed, by to and as are the of of whereas in the is as if the the wild-type of other in the to in the second of in the any When biology the of the and years it out that Ed's was in its the mutation the to be in the second and The is the of the to to Lewis The into the of to into Lewis The second of Ed's career in Drosophila in with a in focus from their and to He both his and his out the to are to the of and would like to be called the The is that the which are to and be with to the and of did Ed that he was to a of about how which could in have been from the on During the Ed began to study that as of the we the He also started to the and of by through his analyses of he was to whether the the of whether they He began to study as as to in the One is Ed's of the which he both and Ed a carrying the wild-type of the into an (Fig. of the during results in of in an wild-type (Fig. with this a that to in which it is Ed to the and that the in the of the The was from the an of this results in of in an wild-type wild-type can an in an that Lewis the in Ed that the a of that of other to come into he this to that the would through both and years analyses this to be the was to and Ed's is for the of It years worth of in It is very to read but it is an paper that is worth the In the to that his Ed from in to the in the on the to the for of the work of and in the late out for mutants with the with Ed in the of Ed's is his of can be called genetics and the wild-type of from their a that Ed had very to the for years. In genetics began by the entire and then by wild-type of the In this Ed was to the and the wild-type of for which he had not obtained also a that genetics fall on the of the one can the of the wild-type Ed was to invent genetics over the he had a of that him to the and then it back one at a time (Fig. with a for the in which all like the second (Fig. he was to back wild-type of the and how That him to that there are in the which on one at a time from more more the of any would be by the of the on in it (Fig. Ed's collection of that the him to invent genetics. A of the is at the are of the available are to the as as to the Lewis with from of the entire results in of all and A wild-type is on the genetics of the to and which and Ed's use of analyses of Lewis with from The for Ed that the on from to the and on in the more the of any is by the of all active in that The second has active whereas has all of the Lewis with of In his Ed also up on a which Pam Lewis had in his lab in the of and the (Fig. Ed that it be a of the and In of his in there would be all the would be on in all which the of the is a of the and from carrying two of the and are the in and Lewis A in the of in with the a at Stanford, and Ed (Fig. to the first of a in the the first which the the the and the on the of the issue of Science in which the was in was Ed's (Fig. Ed for the first time saw the that he had and on the from the (Fig. Ed with at the on and Biology of Drosophila, in in in the first of a and the first Lipshitz (2004). [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com]. Ed's the of Science in which the first of the Reprinted with from [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com]. the of the Ed saw the he had during the on the of the first in that is a of which was to the left to the two had after and had been by in by Reprinted with from The analyses that Ed in his had been both right and in that the in the had by there are in the which are by the (Fig. in that of the that Ed had in the are in that the and of of the One could not have this on the of Ed's the of with was to the There are in the of which the of the rest of the that the and of these Lewis [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com]. The analyses after the of the to the in that the are in from to Ed's the of the in the to the of the that they the all the from to and the the of the of and over years It was this and of of Ed's analyses that to the in that it was time to him the (Fig. Ed the in from of in and who the with Ed, can be to Ed's left in the and from the [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com]. in had in Science that in Drosophila for this he the in 1946. and that the of in Drosophila is in to the and over the it was that given to Drosophila in over a the same effects as an all at and One in highlights that he was about the that the of in also be with their in years he on this the and after by the the War was at its Lewis of the in that in in in the the which could be in any to A.H. Sturtevant of Caltech, in his to the of the for the of Science by that is, in of and be as to the I that an in a position of have that there is from of Ed's into the of and was motivated in by the a of of the present that a would be (Lewis,2004). in late to was in his in Science on and Beadle (Fig. then of the Biology at Caltech, was an to the and that Ed be given to the on In the Ed the from the at and and the in (Fig. He then the first of the in to of for the Ed's of of was two was by a high but it was as had Beadle A.H. Sturtevant and Ed in the late during the that Ed's in Science on and was and motivated public on of the California Institute of Technology his analyses of the and in Ed the in from the at and the of in the in use at that Reprinted with from Lewis Ed on to on other that had been to of who have been for and In his was one two of Ed was to that of the that the of with a of at a to in at a which is to that for the also to the of a which will not In the last of the Ed to which the and the on had would only from the which in they had that a of would be before would be Ed out in the that could in an of by the his of one in a to he a in in the from a of one the of that had been by the It is important to that Ed's in into the effects of was not to it was to with which they could then their about whether to with the There were several of Ed's Science The first of the paper because who was a at Caltech, had Ed's with the that was the of which the on In that used Ed's results to that from from the from the to the In the issue of Science in which Ed's paper was the wrote an called Lewis that there is a the of and the of to Lewis it is to from will in any from any in other of for the it is to the of from as a of any of are the at which it will be to the for a more the It is that the are The are us and we not to we assure other after Ed's he was on the by who that is not a in is to his will who are students of be used as an of the and degree of which be if a as high as were saying he is and you are I I the are that he is and I the are that the on biology and of the is Ed was called to before the US on a of later. was that one of a is a year that one can we this and it that to as the of from year from on by Ed, who was to was the same the for the public of which is had been there would have been year from to had the of to be important to an that a of Ed in of a that was in (Fig. an on on which a of Ed in his Caltech The of was by Lewis of In an in Science he that there is a and He a to in if in a figure which the still Reprinted of Ed the effects of of in several over the to years. a had in that is not in In Ed obtained a from the which him to obtain and then carry out a very of for who had at through from to He that their was that of the US white none of the was from enabling Ed to the that was the of the That study was also the first to that of Ed went on in to the of and had been in their for several found that there was a in their from years after of his was Ed The is an from the on the Biological of have his if the of have (Fig. has supported Ed's the effects of of in Ed in that is at a of to are that from to from I and I of Ed's science with of his effects this one in It on from the in The had that the from Ed out in his paper the to the of and from the that was from that the was in one two He went on to how of in be the important of with whether from other In the of the in the of into the of A study years a in who were at the time of and were to have I not to the that Ed is in any a the he is and of One of is from a in of as a US in The was on the of Biological our were the from other (Fig. Ed up in more which for years he from for the Biology were after by his (Fig. Ed at a in of the author's as a US in May The was in the of the of Biological at Ed, the the a with the and subsequently a with of the Biology at the a with Ed and of the Drosophila at the Ed, [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com]. Ed in two of his from the late After by After by Reprinted from Lipshitz (2004). [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com]. Ed and Pam are at and on other being to by all present as a One in one of Ed's is of Pam at the in where they had been to the of the (Fig. Ed with as Pam in at the in of the of that [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com]. Ed is a In his and are in which he and other The he and Pam as they the of their of is Ed a of and that is with and and He is and in both his and his life. In his he has continued the of and that was by Morgan and his a On a for he has as a and Ed of on he was active in the lab before his Figure the last of his lab with of cross Because Ed began his as a graduate student in he over a year during the He is as a and to The last of Ed's lab cross Ed's cross in Drosophila genetics. of

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