Ellsworth (Ed) Grell blessed the Drosophila community through three enduring legacies: as a pioneer of chromosome mechanics, as a primary organizer and synthesizer of genetic knowledge in Drosophila, and as a graceful mentor to those fortunate to have known him personally.

Ed grew up in rural Nebraska, completed his undergraduate studies at Iowa State, and trained with E.B. Lewis as a graduate student at Caltech, insightfully analyzing the role of two interacting genes involved in the degradation of the amino acid lysine in his doctoral thesis.¹ It was also in the Lewis lab that Ed met and then married his lifetime genetics colleague and soulmate Rhoda Frank Grell. They moved to Oak Ridge, Tennessee, where Ed remained until 1985.

Throughout his career, Ed contributed to developing precisely engineered chromosomes central to creating chromosome balancers, targeted translocations, or chromosome fusions.^2-6 He used these sophisticated genomic tools to follow complex genetic traits in insightful and imaginative ways. For example, Ed was a leader in the arena of linking molecular phenotypes to genetic polymorphisms and enzyme dosage effects, using what were then cutting-edge gel electrophoresis techniques.^7,8 Ed also collaborated with Dan Lindsley at the Oak Ridge National Labs to investigate the genetic basis of spermatogenesis, finding that the haploid post-meiotic phase to generate functional spermatids could proceed in the absence of chromosomes.⁹

Ed is perhaps most appreciated by the Drosophila community for his collaborative work with Dan Lindsley in assembling the “Genetic Variations of Drosophila melanogaster,” commonly referred to affectionately as the Red Book (because of its cover’s color). This wonderful reference book by Lindsley and Grell, published in 1968, distilled all phenotypic and genetic information regarding known Drosophila mutants at the time and provided helpful hints such as ranking mutant alleles on an RK scale based on scoring difficulty. Theodosius Dobzhansky succinctly summarized the achievement of this opus in his Science book review:¹⁰

“Here, at last, is the long-awaited compendium and a guide to the study of the genic and chromosomal variations in Drosophila melanogaster described in world literature. The book modestly calls itself a revision of the C.B. Bridges and K.S. Brehme’s The Mutants of Drosophila melanogaster, published almost a quarter of a century ago. In fact, it is a new and monumental work, embodying what must have been a prodigious amount of meticulous and conscientious work on the part of the authors.”

When Oak Ridge closed down its Genetics Section in 1985, Ed relocated to the Bay Area where he became a central member of the research group led by Lily and Yuh Nung Jan at UC San Francisco until his retirement. Ed helped design both small- and large-scale genetic efforts in the Jan lab and patiently mentored those of us new to the field, sharing his many clever tricks of the trade and his deep historical knowledge of the field. He was a gentle and humble gem of a human being and I feel fortunate to have known him as both a close colleague and friend.

Acknowledgements: Thank you to Lily and Yuh Nung Jan, Susan Younger, Kimberly McCall, and other members of the Jan lab who kindly shared their thoughts and memories of Ed. Thanks also to James Birchler for sharing memories of Ed from his Oak Ridge days and to Michael Mislove for his insights and experience growing up with Ed and Rhoda in their warm and intellectually engaging household.

References:

1. Grell EH. Genetics and Biochemistry of “Red Cells” in Drosophila melanogaster. Dissertation (Ph.D.), California Institute of Technology. 1958. Doi: 10.7907/B5AB-S991.
2. Grell RF, Grell EH. The Behavior of Nonhomologous Chromosomal Elements Involved in Nonrandom Assortment in Drosophila Melanogaster. Proc Natl Acad Sci U S A. 1960;46:51-57.
3. Brosseau GE, Nicoletti B, Grell EH, et al. Production of Altered Y Chromosomes Bearing Specific Sections of the X Chromosome in Drosophila. Genetics. 1961;46:339-346.
4. Grell EH. Variations in Preferential Segregation of Chromosome Two in Triploid Females of Drosophila Melanogaster. Genetics. 1961;46:1267-1271.
5. Grell EH. Distributive Pairing of Compound Chromosomes in Females of Drosophila Melanogaster. Genetics. 1963;48:1217-1229.
6. Twardzik DR, Grell EH, Jacobson KB. Mechanism of suppression in Drosophila: a chance in tyrosine transfer RNA. J Mol Biol. 1971;57:231-245.
7. Grell EH, Jacobson KB, Murphy JB. Alcohol Dehydrogenase in Drosophila melanogaster: Isozymes and Genetic Variants. Science. 1965;149:80-82.
8. Grell EH. Genetic Analysis of Aspartate Aminotransferase Isozymes from Hybrids between DROSOPHILA MELANOGASTER and DROSOPHILA SIMULANS and Mutagen-Induced Isozyme Variants. Genetics. 1976;83:753-764.
9. Lindsley DL, Grell EH. Spermiogenesis without chromosomes in Drosophila melanogaster. Genetics. 1969;61:Suppl:69-78.
10. Lindsley DL, Grell EH. Drosophila Handbook: Genetic Variations of Drosophila melanogaster. Science. 1968;162:993.

Dick on sabbatical and my good fortune

Guest post in memory of Richard Lewontin by Thoru Pederson.

When I was a graduate student in the Zoology Department at Syracuse University, a visiting professor took an office across the hall from the lab where I was stationed. He came “across” to me (not a cis-trans test of allelism but just his kind social reach). At the time I had no idea who he was, so I was exempt from fawning.  

After a couple of chats and lunches, we somehow hit it off. When I told him about my research, measuring the DNA contents of cells in Feulgen-stained preparations, he asked me how I was storing and processing my data. “Storing my data?” I replied. He immediately took my notebooks over to the computer center to see how they could process my data on IBM cards sent into a machine the size of a railroad freight car. I was so struck by his kindness and prescience.

During his sabbatical visit, he gave a series of lectures. After one, a question came: “Are you sure your contention of minimal human genetic variation can be reconciled with what is evident around us?”  He offered the following: “On your way home after this seminar, you might see a rough guy on a motorcycle whereas you may be on your way to your study and a glass of sherry. Nothing in your or his DNA has defined these two paths.” It was a powerful comeback and one I can still remember to this day.

Just at the time of his sabbatical, Dick’s two papers with Jack Hubby came out in GENETICS. All measurements of variation had been based on breeding crosses and now there was a new tool. This was an overnight sensation. (Henry Harris at Oxford got this too but published in the Philosophical Transactions—a huge mistake).  

Dick lived to see the molecular basis of human genetic diversity come to the forefront. In our letters, he never surrendered his position. I will always remember him for the warmth and interest he had for me as a student. But I am sorry that he never recanted on his opposition to Ed Wilson, if not on the core merits but as to the way he waged the battle.

About the author:

Thoru Pederson is the Vitold Arnett Professor of Cell Biology in the Department of Biochemistry and Molecular Pharmacology and Associate Vice Provost for Research at the University of Massachusetts Medical School.

Jean Robert David, the last active member of the French generation who significantly contributed to the establishment of the nine species in the Drosophila melanogaster subgroup as a model for evolutionary genetics, passed away on June 19, 2021, aged 90.

Jean dedicated his 70-year academic career to studying Drosophila biology and evolution, starting as an assistant teacher of biology at the University of Lyon in the early 1950s. His master’s thesis, under the supervision of Victor Nigon (1920–2015), a major figure in the history of Caenorhabditis genetics, concerned the effects of temperature and nutrition on the vestigial mutant in D. melanogaster. There, he made the surprising observation that homozygotes developed the vestigial phenotype when reared on fresh medium; but if reared on old medium (i.e., medium used by larvae from a previous generation) their offspring developed normal wings. This influence of nutrition on the genotype-phenotype map fascinated Jean, and his doctoral work analyzed those effects and their possible transgenerational transmission. Jean’s PhD thesis and subsequent research focused on life history traits (e.g., fecundity, developmental time, longevity, body size, etc.). He believed that fitness effects of genotypes in the laboratory were best understood using optimized media that minimized dietary effects. After testing the effects of a plethora of nutritional elements and food conditions, he developed a standard axenic medium that remains among the most widely used. Following a short postdoc in Edinburgh (Scotland) with Forbes W. Robertson (1920-2012) on Drosophila quantitative genetics, Jean returned to Lyon, where he assumed a professorship position. His laboratory studied environmental effects on phenotypic variation, focusing on the physiology of egg production, food intake, and oviposition-site choice.

Although Jean was a passionate naturalist with a keen interest in insects (he described a new beetle species from the Saone valley in 1963 and a new earwig subspecies from Mont Mézenc in 1973), his work on Drosophila in Lyon was limited to a few laboratory strains. An encounter with researchers from Gif-sur-Yvette, in particular Charles Bocquet (1921-1977) and Léonidas Tsacas (1923-2016), profoundly changed his career and research interests. In the late 1960s, Tsacas secured a major grant to study Afrotropical drosophilids. He invited many French scientists to participate, including Jean who spent one month in Gabon in July 1970. This turned out to be a life-changing experience. First, he realized that Afrotropical D. melanogaster differed in multiple morphological, physiological, and behavioral aspects from worldwide, temperate populations. Second, he became aware of the great taxonomic and ecological diversity of Afrotropical species. Both observations tightened Jean’s collaborations with the Gif laboratory. With Bocquet, he demonstrated in a Nature paper the significant genetic and phenotypic variability among natural populations of D. melanogaster and its sibling species D. simulans, a variation that had not been seen when only non-African populations were studied. For morphological traits like body size, the variability showed parallel latitudinal clines between the two species, likely due to convergent adaptation to cold climate. For other traits such as alcohol tolerance, however, only D. melanogaster populations showed clinal variation with Northern flies being more adapted to highly fermenting resources. Temperature and alcohol became the two main environmental factors that Jean would focus on in subsequent experimental studies.

Jean quickly went from one field expedition to another, collecting flies on islands as remote as Guadeloupe, Martinique, La Réunion, Mauritius, and  Seychelles, and also in Singapore and several African countries. He and Tsacas maintained a strong collaboration by sharing the flies they collected. At the beginning of Tsacas’ project, only three species of the melanogaster subgroup were known: D. melanogaster, D. simulans, and D. yakuba, with melanogaster and simulans cosmopolitan and yakuba endemic to Africa. In 1971, Tsacas described a fourth species from Zimbabwe, D. teissieri; and in the same year, his PhD student Daniel Lachaise (1947-2006) discovered a fifth species from Côte d’Ivoire, D. erecta. Crosses between all those species were impossible or very difficult. In 1973, Jean discovered a sixth species, D. mauritiana, from the island of Mauritius. Unlike the previous species, D. mauritiana readily crossed with D. simulans, producing sterile hybrid males but fertile females. Its close relation to both D. simulans and D. melanogaster made it ideal for studying the genetic basis of speciation in the subgroup. In 1975, Tsacas, Lachaise, and Jean made a joint expedition to Cameroon that led to the discovery of a seventh species of the subgroup, D. orena. Those findings established the Afrotropical origin of the melanogaster species subgroup and pioneered the clade as a premier model for evolutionary genetics.

Following Bocquet’s death in 1977, Jean resigned from the University of Lyon and became the director of the Gif laboratory. Under his directorship, the laboratory continued the study of the evolutionary genetics of the melanogaster subgroup. The 1980s saw two main findings. First, through the application of more advanced biochemical and molecular tools, the dispersal routes of D. melanogaster became clearer. In addition to reconfirming the “out-of-Africa” hypothesis, these biochemical markers helped Jean demonstrate that alcohol-resistant D. melanogaster in Congolese breweries were in fact European flies recently introduced to Africa by man. The European-derived Congolese flies showed signals of reproductive isolation from native African flies, representing one of the most intriguing cases of incipient speciation in Drosophila. A second major finding of this period was the discovery of the specialization of D. sechellia, the eighth species of the melanogaster subgroup, on toxic noni fruits on the Seychelles archipelago. Jean had explored those islands in 1977 but did not pay attention to the smelly noni fruits and their hovering drosophilids. Yet, two students from Oxford did, and their collected flies preserved in alcohol were described by Tsacas and Gerhard Bächli in 1981. In the same year, Jean went to the Seychelles, established the first laboratory culture of D. sechellia, and after 10 years of intensive research, he described in a PNAS paper the genetic basis of D. sechellia behavioral and physiological adaptations to noni.

In 1997, Jean retired but continued to work as an emeritus researcher at Gif. Early in this period, he revisited the role of the environment on phenotypic variation but now within a broader evolutionary perspective, comparing the shape of thermal reaction norms for many morphometrical and physiological traits, such as cold tolerance, at both intra- and interspecific scales. This research program produced a wealth of data documenting genetic variability in phenotypic plasticity, a now-popular topic that was then understudied. It required maintaining a large collection of strains that Jean carefully preserved and enriched through continuous fieldwork. His meticulous observation and isolation of the slight phenotypic abnormalities led him to collaborate with several young talented evo-devo researchers unraveling the genetic basis of natural variants. For example, the observation of supernumerary macrochaetes on the back of some D. melanogaster from Morocco led to the identification of a new repressor of the achaete and scute genes. Similarly, describing the loss of a pair of genital bristles between D. yakuba and D. santomea (the ninth species of the melanogaster subgroup, discovered by Lachaise in 1998) identified a single regulatory mutation of scute with a strong pleiotropic effect on multiple sensory targets involved in copulation. In both cases, Jean’s openness and willingness to share his observations led to new discoveries. 

Jean also had an interest in investigating species not previously studied. He pioneered analyses of the Afrotropical genus Zaprionus and closely monitored the invasion of Brazil by Z. indianus. He created strong ties with Brazilian drosophilists and was frequently an invited guest at the annual Drosophila conference in Brazil. Following Lachaise’s unexpected death in 2006, Jean preserved the laboratory’s tradition of exploring the African fauna, mostly focusing on Madagascar and its surrounding islands. On the island of Mayotte in 2013, at age 82, he made the last major taxonomic discovery in the melanogaster subgroup. He found a subspecies of D. yakuba specializing on the toxic fruits of noni, like D. sechellia. Again, Jean shared his discovery with young population genomicists. This led to detecting a signal of parallel genetic changes between two distantly related species specializing on the same host.

Jean kept working on his experiments in the laboratory at Gif until he fell ill in October 2020. Reflecting on his long career during his terminal illness, he remembered how reading Jean-Henri Fabre’s (1823-1915) 10-volume Souvenirs entomologiques when he was a teenager influenced his view of the living world. Like Fabre, Jean was an “inimitable observer” who mixed a passion for fieldwork with rigorous laboratory experimentation. He was a prolific writer, publishing more than 400 papers, mostly on drosophilids. His style emphasized clarity, using simple words and statistical tests rather than obscure and complex ones, and avoided unwarranted generalizations and fashionable hypotheses. An avid traveler, talented cook (his fruit tarts and chimney-made grills were unrivalled), warm host, and empathic listener, Jean combined an intellectual aura with a genuinely down-to-earth personality—a rare and wonderful combination.

Jean is survived by his beloved wife, eminent developmental biologist Nicole Le Douarin, two daughters, three grandchildren, and two great-grandchildren.

About the Authors:

Amir Yassin – Laboratoire Evolution, Génomes, Comportement et Ecologie (EGCE), CNRS, IRD, Université Paris-Saclay, Gif-sur-Yvette, France

Patricia Gibert – Laboratoire de Biométrie et Biologie Evolutive (LBBE), CNRS, Université Lyon 1, Villeurbanne, France

Pierre Capy – Laboratoire Evolution, Génomes, Comportement et Ecologie (EGCE), CNRS, IRD, Université Paris-Saclay, Gif-sur-Yvette, France

Photo taken by Angelika’s daughter, Theresa Weis

When you’ve been in science long enough, you become part of a certain generation: people with whom you were roommates at Gordon conferences as postdocs, discussed your latest data and shiny ideas during session breaks as a young faculty, and looked forward to seeing at meetings as established scientists. Angelika was part of my generation; we met as postdocs and remained friends long after our scientific interests diverged. Over the years we talked a lot about science but also about kids and dogs and gardening. One doesn’t expect to write a tribute for someone from their own generation—at least not just yet.

Angelika was born in 1967 in Vienna, Austria. She did her PhD with Kim Nasmyth at the Research Institute of Molecular Pathology, also in Vienna, studying the regulation of the cell cycle in budding yeast. She then came to the US to do her postdoc at MIT. After a brief period with Ruth Lehman, who moved to New York, Angelika became a Whitehead Fellow, returning to the study of cell cycle regulation and, in particular, the spatial and temporal regulation of events associated with mitotic exit. Angelika’s lab was one of the first to demonstrate that the nucleolus can serve as a reservoir for regulatory proteins that act elsewhere. Angelika then joined the Biology Department at MIT, where she ultimately became the Kathleen and Curtis Marble Professor in Cancer Research at the Koch Institute for Integrative Cancer Research at MIT and a Howard Hughes investigator. She expanded her work from yeast to mammalian systems, exploring the consequences of aneuploidy, which is when a cell has a wrong complement of chromosomes, a condition often associated with cancer. Angelika’s lab showed that different types of aneuploidy are associated with a common signature of cell stress and altered metabolism. Her lab further showed that this, in turn, alters cell properties in ways that both promote tumor progression and may increase resistance to chemotherapy. Beyond her studies on aneuploidy, Angelika was a true believer in curiosity-driven basic research, which she practiced to tremendous success in studies on cell size and density, mitochondrial surveillance pathways, the role of long non-coding RNA in yeast sporulation, the role of meiosis in cell rejuvenation, and more.

Angelika’s groundbreaking discoveries led to many awards, too numerous to list here. They include the Breakthrough Prize in Life Sciences, which “honors transformative advances toward understanding living systems and extending human life”¹ and the Vilcek Prize in Biomedical Science, given to “immigrants who have made lasting contributions to American society.”² Angelika also won the Genetics Society of America Medal, which “honors an individual member of the Society for outstanding contributions to the field of genetics.”³ She was elected to the National Academy of Science and the American Academy of Arts and Sciences. Her devotion to science extended well beyond research, and she served on many boards and committees, including the Genetic Society of America’s Board of Directors. She was a fierce supporter of increasing the representation of women in science and an advocate for work/life balance. She did work relentlessly, and she always had her laptop with her—while watching TV, cooking dinner, or attending her daughters’ soccer games. She never stopped thinking about science. But at the same time Angelika would regularly leave work early to spend time with her two daughters, Theresa and Clara, and to allow her husband to pursue his career. She had a beautiful herb and vegetable garden that she tended to herself, and she loved orchids and cooking Austrian dishes. She married her high school sweetheart, Johannes, and they remained devoted to each other ever since. Whatever balance she found must have worked because her daughters are truly remarkable.

Angelika was many things to many different people. She was a brilliant scientist, a trail blazer. She was rigorous in her science and demanded the same from others. She was critical but constructive, and her presence at meetings elevated the level of discussion. In her own work, Angelika would seize on an observation, imagine its underlying mechanism, set out to investigate it, and, more often than not, figure it out, which brought her a tremendous amount of joy. I often wondered what made her such a successful scientist; she was definitely bright, but most scientists are bright. What set Angelika apart, I think, was her instinct, which rarely failed her; her conviction, which prompted her to fearlessly pursue a biological question, regardless of the unknown; and her unrelenting belief in her people. It helped, of course, that she attracted brilliant students and postdocs. Still, Angelika’s boundless and infectious enthusiasm inspired people to believe in themselves. Angelika was a dedicated mentor and avid cheerleader to her trainees. She wrote the most amazing letters of recommendation for the people in her lab, and they, in turn, did not disappoint. Angelika made a lasting mark on science not only through her scientific discoveries but also through the scores of talented and well-trained scientists, many of whom are women, who came from her lab.

And, of course, Angelika was immensely entertaining. She was funny without even trying to be so. We were amazed by her boldness, her willingness to share funny personal anecdotes with near strangers. She could be both naïve and extremely savvy. Knowing Angelika was knowing where all the diet coke vending machines were at every meeting venue you ever attended together. It was knowing that when she said, “May I say something?” it was really a rhetorical question. We admired her openness, her seize-the-day approach to life, her joy and sense of wonderment when learning something new, science-related, or otherwise. Angelika was larger than life; she was a force. You would hear her laugh from across the room. Angelika spoke her mind and made us better for it. But the thing that made Angelika so lovable was that even with all her accomplishments, awards, and famousness, she always treated people as her equal—grateful for their advice, time, and friendship. She had many friends, and she loved them all fiercely.

Angelika passed away in October 2020 at the age of 53 after a two-year battle with cancer. She will be sorely missed.

Written by Orna Cohen-Fix

NIH, NIDDK

¹https://breakthroughprize.org/

²https://vilcek.org/prizes/vilcek-prizes/

³https://genetics-gsa.org/awards/genetics-society-of-america-medal/

Lea Kanner Bleyman
November 9, 1936–November 6, 2020

GSA member Lea Kanner Bleyman died on November 6, 2020, three days short of her 84th birthday. Lea was a Professor Emerita in the Department of Natural Sciences at Baruch College and a past-President of the International Society of Protozoologists (now the International Society of Protistologists). 

Lea was born in Halle, Germany on November 9, 1936. After escaping with her family to France in 1939, just before World War II, Lea spent time sheltered in OSE homes (for Jewish orphans) before a stint as a hidden child, living in a convent and on a farm to protect her from Nazi terror. Post-war, reunited with her parents and two older sisters, she started a new life in New York City in 1946. For her Bachelor’s degree, Lea attended Brandeis University on a full scholarship. She earned a PhD in Genetics at Indiana University in 1966. As a professor, Lea delighted in instructing her students, which she far preferred to her year serving as Chair of the Department. 

Lea K. Bleyman and her daughter, Anne, on a trip together in Iceland.

Away from the classroom or the lab, Lea was known for her vivacity. She was a voracious reader, and she subscribed to the New York Times primarily for the crosswords and other puzzles, which she completed in ink. Lea loved opera, cats, theater, college sports, sharing learning experiences with new friends in Shakespeare and opera classes, and getting letters to the editor published in the New York Times.

She is survived by her daughter, Anne Bleyman; her husband, David Minn; sister Eve Kugler; nieces and nephews.

We are sad to report that Dr. Karl Gordon Lark died on April 10^th, 2020 from an aggressive form of prostate cancer. Karl Gordon Lark was born on December 13^th, 1930 in Lafayette, Indiana to Karl and Betty Lark-Horovitz. His father was a physicist with an interest in biology. Gordon entered the University of Chicago in 1945. Following a course in physical chemistry in 1948, his professor, Leo Szilard, recommended the newly emerging field of molecular biology and, as a good introduction, Gordon took the phage course at Cold Spring Harbor taught by Marc Adams. Subsequently, Gordon enrolled at New York University to obtain his PhD with Marc as his mentor. He became a member of the “Phage Group”  of scientists under the guidance of Max Delbrück, using bacterial viruses as a model organism for genetics. This group included the likes of not only Delbrück, but also Luria, Hershey, Meselson, Stahl, Watson and Crick. Gordon obtained his PhD in 1952 and began a lifelong career in molecular biology with his companion and wife, Cynthia Thompson. Gordon Lark emerged as a remarkable scientist not only with an illustrious career in the molecular biology of bacteria and bacteriophages, but he then ventured into new territories including soybean and dog genetics. In each of these fields a hallmark of Gordon’s pursuits were clarity, passion and creativity.  How he managed, with only a modest-sized laboratory, to significantly contribute to such a diverse set of fields was, and still is, truly amazing.

In 1970 Gordon Lark was recruited to the University of Utah to assemble a new Department of Biology that would include all of the biological sciences, from molecular biology to ecology and evolutionary biology. In the first six years he hired seventeen new tenure-track faculty. His energy was boundless, his formula simple. He looked for excellence, creativity and synergy. The latter he interpreted as having passion for science and the need to share that passion, particularly with the next generation of scientists. In a few years, Gordon’s vision, tenacity and impeccable taste established a world class Biology department that was particularly well-recognized for its excellence in genetics. New recruits learned so much from Gordon, who taught by example. His rigor, diligence and attention to detail were unparalled.  He set very high bars for himself and his science, with those around him sometimes struggling to keep up. But, perhaps surprisingly given his boundless energy, Gordon was in some ways a very patient man. Before every critical faculty vote Gordon would take time to personally discuss this issue with every member of the department. When the vote came, there were no surprises.

The dog genetics community owes a special thanks to Gordon for his seminal contributions. His PNAS paper in 2002 with Kevin Chase revolutionized the field, demonstrating for the first time that seemingly complex morphologic traits were controlled by a small number of genes, thus nominating the dog as a genetic system for the identification of genes controlling breed-specific differences in morphology and behavior. That paper also provided a key impetus to select the dog as a mammal for early whole genome sequencing in 2005, which in turn allowed canine genetics to blossom as a system of interest to medical geneticists, anthropologists, behavioral scientists and physiologists.

Gordon was unique in his ability to make connections where no one else saw them. He frequently drew parallels between his soybean and dog work, pointing out that interactions between quantitative trait loci in soybeans were key for understanding phenotypic variation. He argued, correctly, that the same must be true in dogs, likely explaining some of the nuanced changes observed between breeds.  The “Georgie Project,” an effort to fully sample large numbers of living Portuguese Water dogs is just one example that can be pointed to when a trainee says “…I can’t possibly…,” reminding them that if Gordon could do it, they can at least try.

Gordon Lark created a space of trust and synergy, rather than competition, a place that inspired, was rigorous but also joyful. The surrounding environment in Utah is magnificent, and its splendor matched the environment Gordon built in his department. It is doubtful that any achievements which we have been fortunate to accrue could have been attained in any other environment. Utah was and is a place where people like Gordon could think big, and then bigger still, with unfettered creativity. Those of us living in other places found reasons to visit Gordon frequently, believing that our collaborations with him would somehow make us heirs to his success.

Our final mental pictures of Gordon are not of him at the bench, but of him with Mopsa, his all-time favorite dog, sitting beside him in his messy office as Gordon talked with unbridled enthusiasm about the achievements of his family, and his love of music, literature, and history. We will all miss Gordon. In these very strange times of social distancing, it is difficult to grieve together. However, such grieving, in Gordon’s eyes, would likely be displaced. What would please Gordon most is for us as scientists to continue our endless quests for the joy of doing science and, most of all, to share that joy with the next generation of scientists.

Mario R. Capecchi

University of Utah School of Medicine

Elaine A. Ostrander

National Human Genome Research Institute/NIH

Bruce Stewart Baker—the geneticist whose work uncovered molecular mechanisms of Drosophila sex determination and dosage compensation—died unexpectedly on July 1, 2018. He was 72.

Bruce Baker was born on Dec 20, 1945 to William K. (Bill) Baker—also a renowned Drosophila geneticist —and Margaret I. Stewart in Swananoa, NC, the site of the closest army hospital to where his father was stationed during WWII.  Bill and his wife hosted what turned out to be the inaugural Annual Drosophila Research Conference (ADRC) in their home in 1958—with a total of four Drosophila biologists sharing their findings with each other and with a small group of geneticists studying other organisms. The ADRC currently includes over 1500 participants.

Bruce attended high school in Chicago, but left before finishing because he was accepted by Reed College in Portland, Oregon, one of the few places in the country that did not require a high school diploma if one’s SAT scores were high enough. After graduating from Reed in 1966—where he apparently spent quite a lot of time playing poker—Bruce went to the University of Washington where he did his graduate work with Larry Sandler, generating and characterizing X-linked meiotic mutations and the meiotic mutant pal. Bruce received his PhD in 1971 and spent another year doing research in the Sandler Lab before moving to Madison, Wisconsin to do two years of postdoctoral training with James F. Crow. Bruce then spent two years as an Assistant Professor in the Zoology Department at the University of North Carolina, Chapel Hill, before moving to the Biology Department at the University of California, San Diego (UCSD). At UCSD, Bruce rose through the ranks of Assistant, Associate, and Full Professor, sharing his faculty position with his partner at the time, Adelaide T.C. Carpenter. In 1986, Bruce took a position as a Professor in the Biology Department at Stanford University, where he remained for the next 22 years. In 2008, he moved to the Janelia Research Campus of the Howard Hughes Medical Institute, from which he retired in 2016.

Bruce received many accolades throughout his illustrious career. He received the 1991 Genetics Society of America Medal. He shared the National Academy of Sciences Award in Molecular Biology with Professor Thomas Cline in 1992 for their creative use of genetics and molecular biology to uncover how sex is determined in Drosophila. Their work demonstrated that assessing the number of X chromosomes (relative to autosomes) can initiate an RNA splicing cascade that controls all aspects of somatic sexual dimorphism, from external morphologies to sex-specific mating behaviors. Bruce was elected to the National Academy of Sciences in 1993. He was vice-president and then president of the Genetics Society of America in 1993 and 1994. From 2001 until his death, Bruce was the Dr. Morris Herzstein Professor (and then, Professor Emeritus) of Biological Sciences at Stanford.

In his career, Bruce published over 150 papers, most focused on the cellular and genetic mechanisms of how sex—and dosage compensation—is achieved in Drosophila. It was at UCSD that Bruce began his pioneering work uncovering the molecular mechanisms of Drosophila sex determination and dosage compensation. In a seminal paper published in 1980 “Sex and the single cell. I. On the action of major loci affecting sex determination in Drosophila melanogaster”, Bruce first described the phenotypes associated with mutants in each of the major loci affecting sex determination [alone, in combination, and in mosaic clones], revealing that sex in flies is determined at the level of individual cells (i.e. is cell autonomous) and that the genes required for achieving normal sexual morphologies continue to be required in adults to regulate sex-specific courtship behaviors. In the decade that followed, Bruce and postdocs in his lab cloned and molecularly characterized many of the major players regulating sex determination:  transformer (tra), transformer-2 (tra-2) and doublesex (dsx). The characterization of these genes led to the surprising finding that sex determination in flies is largely controlled at the level of mRNA splicing, culminating in the production of sex-specific forms of the Dsx transcription factor—whose orthologues function in sex determination in all higher animals, including humans. Through the early 1990’s, Bruce and his team continued to molecularly characterize factors controlling Drosophila sex, including Fruitless, a transcription factor that is also regulated by Tra/Tra-2-dependent splicing and plays a major role in male-specific courtship behaviors. During this time, his group also molecularly characterized several regulators of dosage compensation (male-specific lethal genes) that were shown to be components of a large protein-RNA complex that binds to genes on the single X chromosome in males and increases levels of expression of X-linked genes to equal the level generated from the two copies of these genes in females, which are chromosomally XX. In the two decades that followed, Bruce and his collaborators expanded their studies into the evolution of sex determination, the neural circuitry of sex-specific behaviors, and how the sex determination cascade is integrated with input from other developmental hierarchies to control sexually dimorphic development. Their work revealed key downstream targets and uncovered the neuronal wiring underlying the different behaviors of males and females. Bruce also maintained an interest in genes that function in both meiosis and mitosis, publishing a number of papers with his long-term friend and collaborator Maurizio Gatti, from the Sapienza University of Rome. At every stage in his career, Bruce immediately embraced emerging technologies to answer questions about how sexual morphology and behavior are controlled at the molecular and cellular level.

In the early days of his career, Bruce spent nearly all his waking hours in the lab—seven days a week, nearly 364 days a year (he took Christmas off), maintaining his focus with serial cups of coffee shipped in from Seattle and brewed in an Erlenmeyer flask warmed over a Bunsen burner. Maurizio Gatti remembers returning to the lab with Bruce after dinner, where they often had long late-night conversations not only about science but also history, politics, trekking in wild places, human relationships, personal feelings, and life in general—conversations that Maurizio always found interesting and pleasant because Bruce had not only a wide and deep knowledge of the facts, but also a rather peculiar sense of humor. That same sense of humor often helped break up tense situations in the lab, when Bruce would suddenly giggle after giving a long piercing look that made one feel he was reading your mind, remembers a former student. Bruce had enormous respect for the details of science, not only the science in his own lab but also that of his peers; many of his trainees remember spending several weeks of journal club on a single Tom Cline laboratory paper (some remember several weeks on a single figure). It was that attention to detail that led to Bruce’s insightful models for sex determination and dosage compensation that were elegantly simple and explained everyone’s findings.

Bruce loved the outdoors—trekking, skiing, and fishing, even tying his own flies. His rare escapes from the lab involved trekking expeditions in Nepal, the Fairweather Range, and around Denali, Alaska with his early partner Adelaide Carpenter (one August Alaska trip included getting caught in a blizzard, but surviving with only minor frostbite on the tips of some toes). Bruce also enjoyed cooking, with a particular passion for Cajun, Creole, and Thai food. Many of his trainees and collaborators benefitted from the elaborate meals he would produce of each type of cuisine. Something most of his students and colleagues did not appreciate was Bruce’s skill at the computer game “Civilization”; for several of his years at Stanford, Bruce was the top scorer in this game under the pseudonym “King Bruce”. Bruce must have had to find something new to do in the lab late at night after his group grew large enough so that he didn’t feel like he had to work so hard all the time. Bruce was proud of all of his trainees that went on in science, either to labs of their own or to other science-related careers. He was also immensely proud of and devoted to his wife Allison C. Chin, an environmentalist who served in key leadership positions in the Sierra Club, including as President, and who now serves on the governing boards for the Sierra Club Foundation, Women’s Voices for the Earth, and North American Association for Environmental Education (NAAEE)—an organization dedicated to accelerating environmental literacy and civic engagement through education. Their adventures included backpacking on Baffin Island, rafting the Aichilik River in the Arctic National Wildlife Refuge, and exploring the Antarctic Peninsula. To all who knew him, Bruce was a quiet man with unparalleled scientific rigor, who was willing to stand up to authority for what he believed. He was an inspiring teacher, practicing the Socratic method in the classroom and in the laboratory. Indeed, while he continued to carry out his own experiments even through very late stages in his career, Bruce was also delighted to take time at the bench to teach his trainees—for example, showing them how to “read” the giant salivary gland polytene chromosomes, pointing out all the morphological landmarks, such as the “ballet skirt”, the “onion”, the “duck’s head” and the “goose neck”.  For the many of us who still love and do science, we carry with us both Bruce’s high scientific standards, his love of hands-on teaching at the bench, and his unparalleled enthusiasm for discovery.

Guest post by Deborah Andrew with the help of many friends and colleagues of Bruce Baker.

Guest post by Nina Wolff pays tribute to long-standing GSA member Margaret Lieb.

Margaret (Peggy) Lieb died on March 8, 2018 in South Pasadena, California at the age of 94. After attending schools in New Rochelle, NY, she graduated magna cum laude from Smith College, and subsequently studied with  H.J. Muller at Indiana University and with Francis Ryan at Columbia University, where she received her PhD degree. Following postdoctoral studies at Caltech in the laboratory of Max Delbruck, and in Paris at the Pasteur and Radium Institutes, Lieb taught at Brandeis University and then moved to the Medical School of the University of Southern California where she continued her research and teaching for 45 years. After her retirement, she continued to be active as an Emerita member of the faculty, and as a garden docent at the Huntington Museum and Botanical Garden.

While at Caltech, Lieb published one of the first studies of phage lambda, and subsequently isolated and characterized a large number of mutations in the repressor gene of the phage. Her studies of lysogenization indicated that the active repressor was a dimer, a conclusion later confirmed by biochemical studies in other laboratories. While mapping mutations in the lambda repressor gene, she observed that excess recombination (negative interference) was associated with mutations arising from the deamination of 5-methylcytosine. This led to the identification of a novel mismatch repair gene (vsr) in E. coli – a gene that is adjacent to the gene for cytosine methylase. The Vsr function reduces the probability of mutations that occur due to spontaneous deamination of 5meC. Although genes related to vsr appear to be limited to bacteria, the search for genes like vsr in eukaryotes, where 5-methycytosine has important regulatory functions, has led others to the discovery of additional specific repair activities in higher organisms.

In 1972-1973, Lieb served as Program Directory of the Genetic Biology program of the National Science Foundation. She was elected Chairman of the Virology Division of the ASM in 1975, and served on the editorial boards of Journal of Virology and GENE. She was a Fellow of the American Association for the Advancement of Science (AAAS).

Peggy Lieb maintained an active interest in the research of her colleagues, and will also be missed by the students and post-doctoral fellows who spent time in her lab. Her high standards of performance in the classroom and in the lab were challenging and also appreciated by those who knew her.

Guest post by Thomas Kaufman in memory of Kathy Matthews.

The original Drosophila Stock Center was housed at Cal Tech in Pasadena and was established by Calvin Bridges and Alfred Sturtevant—students of Thomas Hunt Morgan. On Morgan’s and Bridges’ deaths, the responsibility for the collection fell to Sturtevant, who subsequently passed it to his student Edward Lewis. In the early 1980s, this collection contained ~1600 stocks, representing mostly classic mutations, chromosome aberrations, and balancers that had been accumulated over the years. The center was supported by the National Science Foundation (NSF), and it distributed stocks to the Drosophila community at no cost.

How did the Cal Tech collection move to Indiana University?

In the early 1980s, the Annual Drosophila Research Conference (ADRC) was a much smaller affair and could be held at more intimate venues. One of the favorites was Asilomar, a conference center on the Monterey peninsula in California. I was standing in line for lunch and chatting with Dan Lindsley when he mentioned that Ed Lewis was retiring—and that Cal Tech didn’t want to continue hosting the stock collection. It occurred to me that my institution, Indiana University, with its excellent fly facilities and low costs, might be a good place to move the collection. I proposed doing so, and Dan agreed to talk to Ed and help with the move.

My volunteering was not entirely altruistic. At that time, there was a young postdoc in my lab named Kathy Matthews. Kathy had come from the University of Texas where she had gotten her Ph.D. working in the lab of Yuichiro Hiraizumi on the population dynamics of transposons. She was an accomplished geneticist and adept at genetic manipulations. Her work in my lab focused on the genetics, molecular, and developmental biology of the α-tubulin gene family in D. melanogaster. The contrast between her thesis work with her postdoc project illustrated not only her wide-ranging interests but her technical bravery as well. An example of her technical fearlessness lay in her developing 2D gel technology in the lab. In the early 80s, there were no kits, and one could not simply order a fancy gel apparatus out of a catalogue. Kathy built her own 2D gel boxes and proceeded to produce some of the most beautiful 2D gels and westerns I have ever seen. Additionally, she designed and expressed unique peptides from the conserved α-tubulin proteins, allowing her to raise isoform specific antibodies to study their different patterns of accumulation.

I knew that Kathy was not particularly excited about the prospect of a traditional academic position that involved teaching and committee duties. She simply liked doing science without these other distractions. She was smart, knew flies and genetics, and was very well organized. I thought, if she would do it, she would make an excellent manager for the stock collection. Ed Lewis had always had such a person, and it seemed a good idea to continue with his model. When I returned from California, I told her of Cal Tech’s lack of interest in keeping the collection and proposed that we move it to IU—and that she be the collection manager. She agreed, and we began the process of getting the NSF grant transferred to IU by writing a new proposal and making plans to have the flies mailed to Bloomington, thus establishing the Bloomington Drosophila Stock Center (BDSC).

The BDSC collection now contains nearly 68,000 stocks, and the center distributed 218,429 samples worldwide in 2017. The staff has now grown to five senior scientists, four research associates, and fifty part- and full-time stock keepers. Kathy oversaw all scientific and administrative functions and skillfully guided the center as it was supported first by NSF funding, later by a cooperative agreement between the NSF and NIH involving the introduction of user fees, and currently by NIH subsidy.

The Birth of FlyBase

At the 30^th ADRC in New Orleans in April 1989, Dan Lindsley approached Kathy to tell her that the most recent “Red Book” (a compendium of all Drosophila genes, mutations and chromosomal aberrations) was to be his last, and a new way to collect and summarize genetic and genomic information on flies had to be found. He knew the BDSC had its stock lists available at the IU BioArchive, and he wondered if that mechanism could be parlayed to provide Red Book-like material. An informal workshop on Drosophila databases was held at the meeting, with a more in-depth workshop occurring under the auspices of the National Center for Human Genome Research (NCHGR) in 1990 in Bethesda. This meeting concluded with a group of volunteers agreeing to work together on an integrated Drosophila database; Kathy, of course, was one of the volunteers, and work on FlyBase began.

Much like the BDSC, FlyBase has grown enormously both in content and scope. It evolved from a simple relational database of stock data with flat files provided via ftp into an interactive web-based database that provides query tools to access the data. With the addition of newer tools and larger data sets, FlyBase has now evolved into the sophisticated research instrument and knowledgebase that we know today. Kathy was a strong guiding force through much of this development.

A good deal of these two powerful resources would not have been possible without the active, instrumental, knowledgeable, and selfless involvement of Kathy Matthews. Her contributions to the Drosophila research community have been numerous and exceedingly important. We could not do things the way we do them today without her hard work, dedication, and foresight. She was an enormous gift to our community and she will be missed. We are the better for her all too short presence amongst us, and we are the less for her passing.

GSA was saddened to learn about the passing of William Gelbart, a long-time member of the Society, former member of the GSA Board of Directors, former editor for GENETICS, and the 2010 recipient of GSA’s George W. Beadle Award for contributions to the community of genetics researchers.

Bill Gelbart (Photo courtesy Susan Russo)

Bill was professor of molecular and cellular biology at Harvard University. His laboratory focused on understanding the molecular basis of pattern formation in higher animals, focusing on cell-cell signaling related to the decapentaplegic pathway.

Beyond his many research contributions, Bill is well respected for his leadership in the Drosophila community. He has served as Principal Investigator for FlyBase since the project was initiated in 1991. He was also a trusted advisor to many other community efforts—including WormBase, Zebrafish Information Network, The Arabidopsis Information Resource, and many others—and served on the National Advisory Council for the National Human Genome Research Institute.

Ken Burtis and Brian Oliver demonstrate the love of the community for Bill Gelbart. Photo credit: Adam Fagen.

Bill earned his BS in biology from Brooklyn College in 1966 and his PhD in genetics from the University of Wisconsin in 1971. He did postdoctoral work with Ed Lewis at Caltech and Art Chovnick at the University of Connecticut before moving to Harvard in 1976. In addition to his research and teaching during his tenure at Harvard, Bill served as the Head Tutor for the undergraduate concentration in biology and as program director for an interdepartmental predoctoral training program in genetics and genomics.

Bill is survived by his devoted wife, Susan;  his loving daughters Marnie, Courtney, and Jennifer;  his adoring grandchildren Delilah, Theodore, and Amelia; a brother and sister-in-law;  many nieces, nephews and dear friends.   A memorial service honoring Bill will take place at a later date.The family has requested that in in lieu of flowers, contributions can be made to the The Greater Boston Food Bank or WGBH (Boston’s PBS station). Those who wish may also donate to GSA in Bill’s memory.

Additional Information:

• Thomas C. Kaufman, “The 2010 George W. Beadle Medal: William M. Gelbart,” GENETICS 184: 871-872. doi:10.1534/genetics.110.114348.