Julian E. Davies, Ph.D. Professor Emeritus, University of British Columbia [January 9, 1932 – February 2, 2025]

We express our deepest sadness at the passing of Professor Julian E. Davies, our most esteemed mentor and beloved friend of microbiologists worldwide, on February 2, 2025 near Vancouver, British Columbia.

Julian was born in Wales in January 1932, received his Ph.D. in organic chemistry from Nottingham University in 1956, and did postdoctoral work in natural product chemistry at Columbia University in New York and the University of Wisconsin. He was a lecturer at Manchester College of Science & Technology in 1959, a research associate with Bernard Davis at Harvard Medical School from 1962 to 1965 and with Francois Jacob at the Institut Pasteur from 1965 to 1967. In 1968 he established his own laboratory in the Department of Biochemistry at the University of Wisconsin-Madison (UW).

Julian published an invited autobiography entitled “Gathering No Moss” to Annual Review of Microbiology in 2003. We will defer many details of his research results and his friendships to that autobiography.

Here six of the scientists who worked with Julian during the 1970s describe our fond personal memories of our times with him, and his three grown children describe their wonderful family life in Madison, Geneva, Paris, and Vancouver as further illustration of his creative, energetic and joyful character.

Overview by Patrice Courvalin

Throughout Julian’s career, the research in his group reflected aspects of his personality: notably great curiosity, creativity, energy, and sociability. This research was varied, often collaborative, and, most importantly, innovative. As a great admirer of Louis Pasteur, Julian sought to ensure that his fundamental research would have translational and practical applications.

His research programs over the years evolved from his first training in chemistry to microbiology, and molecular biology and genetics. Themes of his research included the discovery and mode of action of antibiotics, their biosynthesis, ribosome structure and function, the origins, evolution and spread of antibiotic resistance among often unrelated microbial taxa of antibiotic resistance: mobile DNAs (plasmids, transposons, integrons); analyses of antibiotic-producing microorganisms, by genetic, biochemical, and ultimately metagenomic methods, the likely environmental roles of these and other natural products, including cell signaling and microbial population control when at the low (sub-inhibitory) concentrations as in nature; and the extent and potential applications of microbial diversity.

In early experiments (1964), Julian showed that streptomycin blocked translation of bacterial mRNAs by binding to 30S subunits of bacterial ribosomes, that resistance due to mutation in laboratory Escherichia coli was due to a change in the 30S ribosomal subunit, and that streptomycin and several other ribosome-targeting drugs could cause errors in mRNA translation (amino acid substitution). Left unsettled was how streptomycin also damages membranes of susceptible cells.

Soon thereafter Julian, with his new group at UW, showed that resistance in many clinical isolates to the antibiotics streptomycin, and the neomycin and gentamicin groups usually resulted from specific enzymatic modification or inactivation of the drug (variously by adenylation, phosphorylation or acetylation), not by changes in the ribosomal targets of their action.

In 1973 Julian and his first Ph.D. student, Raoul Benveniste discovered antibiotic modifying enzymes in extracts of antibiotic-producing bacteria (e.g., Streptomyces) whose activity profiles matched those from drug-resistant pathogens. Hence, the irony that the very sources of such life-saving drugs also might have been the ancestral sources of the resistance genes that increasingly compromise their clinical usefulness. This hypothesis has received strong support over the years by findings (i) of broad range R (resistance) factor plasmids, transposons and other mobile DNAs that facilitate gene transfer among unrelated taxa, (ii) that genes cloned from antibiotic producers transferred into other strains in lab experiments do indeed confer expected resistance phenotypes, and (iii) in 1993, of DNAs that can confer resistance can be found in commercial antibiotic preparations.

Other valuable research during this period included the isolation of restriction enzymes PstI and KpnI, that were useful for physical mapping and recombinant DNA cloning; and a finding that the npt gene of resistance transposon Tn5 (which Julian co-discovered in Geneva; see below), also conferred resistance to G418, an amionglycoside toxic to eukaryotes. This led to the widespread use of G418 and this npt gene for genetic manipulation and analysis of diverse fungal, plant and animal systems. Also, his was one of the early voices warning that the overuse and careless use of antibiotics would select resistant bacteria, making the antibiotics less and less effective.

Practical applications of his basic research were important to Julian. This is illustrated by use of aminoglycoside-modifying enzymes to quantify aminoglycoside serum concentrations, genetic strategies for hyper-producing antibiotics, discovery of new useful restriction endonucleases, and construction of shuttle vector plasmids for Actinomycetes. The appeal of practical applications also led to his move from UW to Geneva in 1980, to become Research Director of the Swiss operation of Biogen, a pioneering international biotechnology company. He returned to Paris in 1985 to become head of the Laboratoire de Génie Microbiologique at the Institut Pasteur. There he began studies of mobile DNA elements and selectable markers in Mycobacterium smegmatis, a non-pathogenic and fast-growing relative of M. tuberculosis, with important potential for vaccine development and analysis of M. tuberculosis virulence genes.

Julian returned to university life in 1992, to become Chair of the Department of Microbiology at the University of British Columbia. There he started the TerraGen Discovery company in 1996, with the goal of finding new antibiotics and other useful compounds in environmental samples. Its core platform entailed encapsulation of individual spores and cells from natural product-producing microbes collected from environmental samples in gelatin beads for growth and screening for antibiotic production. TerraGen was also a pioneer in the isolation of environmental metagenomes and the cloning of entire antibiotic biosynthetic gene clusters decades before this was routine in other labs.

He became interested in effects of antibiotics at the sub-inhibitory concentrations at which they would occur in nature. His found that low levels of these “antimicrobials” change transcription patterns and phenotypes, including virulence traits and biofilm production, of both the producing and other species. These findings have contributed to our understanding of selective pressures that may have promoted the emergence of antibiotic resistance.

Endlessly fascinated by the diversity of specialized metabolites that microbes produce, he developed the concept of the ‘parvome’, defined as the collection of bioactive small molecules produced by bacteria. [J Antibiot, 2017;70(4):339–346. https://doi.org/10.1038/ja.2017.14].

In his later years he studied antimicrobial activities of natural clays, which were well known to Canadian First Nation and other indigenous and ancient peoples to have important healing properties. Focusing on clays from the coastal British Columbian Kisameet Bay region, he and students showed that clay suspensions and clay extracts have complex, potent antimicrobial activity against numerous bacterial and fungal pathogens including ones resistant to multiple traditional antibiotics.

Julian received many honors and recognitions over the years, including election as Fellow of the Royal Society of London and of Canada, member of the American Academy of Microbiology, and Presidency of the American Society for Microbiology (ASM), Presidency of the International Union of Microbiological Societies (IUMS), and International Member of the US National Academy of Sciences (primary section, Microbial Biology; secondary section Environmental Sciences and Ecology).

Memories of Douglas E. Berg

My time with Julian was one of the best periods of my life, professionally and personally. It actually started in 1973 with my reading of his just-published PNAS paper reporting aminoglycoside inactivating enzymes in Actinomycetes whose activity profiles matched those responsible for antibacterial resistance (ABR) in clinical isolates. The evolutionary implications were intriguing — that the sources of these marvelous drugs also might be ancestral sources of ABR genes that increasingly negated such drug’s clinical value. Julian came on sabbatical a year later to the University of Geneva’s Département de Biologie Moléculaire (UniGE DBM). I had been at DBM in Lucien Caro’s group for several years, studying “dv” plasmids, which arise from bacteriophage lambda by in vivo deletion of phage genes not needed for controlled autonomous DNA replication. On meeting Julian I began dreaming of new ways to advance my dv studies: in particular, a 1970s style in vivo cloning strategy (this was before in vitro recombinant DNA cloning protocols and reagents had become available). We imagined that lambda would pick up ABR genes by a phage specific process: inserting nearby during lysogenization, and then excising aberrantly upon induction of lytic phage development to generate transducing phage containing bacterial gene substitutions adjacent to the phage ATT site. Although extremely rare, the pickup of E. coli chromosomal genes in this way had been demonstrated by Robert Weisberg at the NIH. As in Wesiberg’s studies, we anticipated that such R factor genes could be selected among the many billions of phage that were produced from a few mL of induced E. coli culture after en masse lysogenization and then prophage induction.

We were delighted when many hundreds of hoped for “lambda-kan” (resistance transducing) phage were obtained (detailed in our 1975 PNAS paper and a 2017 tribute to Julian in this journal [J Antibiot, 2017;70(4):339–346. https://doi.org/10.1038/ja.2016.120]). This result apparently met one of our goals: the “cloning” of ABR genes, actually very easily! However, physical analyses of DNAs several of lambda-kan phage by DBM faculty colleagues Jean-David Rochaix and Bernard Allet showed that the new DNAs were not the expected substitutions. Rather, specific DNA segments had been cleanly inserted into different sites in different lambda-kan genomes. This was a wonderful surprise, as it implied the discovery of an ABR-containing transposable element (“transposon”), a type of mobile DNA element not previously known. Its existence expanded understanding of how ABR genes could spread in bacterial populations. And, transposons marked with an easily selectable ABR gene promised to facilitate further study of the transposition phenomenon, and also to be valuable for bacterial genetic analysis and manipulation. We focused on one kan element, soon to be called Tn5, based on finding that it also could transpose frequently from lambda to the E. coli chromosome, and from there back to phage lambda.

Julian had come to UniGE with no specific plans other than to do and learn good stuff. I was grateful to benefit there from his creativity, sense of adventure and joy in good experiments and life in general. Importantly, also benefiting from his fine mentorship, was Iwona Stroynowski, a brilliant Polish undergraduate majoring in theoretical physics at UniGE who wanted to consider a more biology-focused career. Julian agreed to serve as her mentor, and shared his small lab room with her. From this vantage point she witnessed and shared in the design of our experiments, and excitement of Tn5’s discovery. Importantly, Iwona credits Julian with teaching her to love bacterial genetics and hands-on lab techniques, and directly inspiring her to leave theoretical physics for a career in biology — a Ph.D. at Stanford (Lederberg and Yanofsky labs), a postdoc in immunology, and then a professorship at the University of Texas Southwestern. It was a marvelous time for all: lots of learning and discovery, buoyed by Julian’s creativity, knowledge, curiosity and good humor, and our friendships, collaborations and enjoyment of life in Geneva and environs.

My plans to stay in Europe for the indefinite future changed when Julian offered me a temporary position in his UW lab to continue Tn5 studies. I moved there soon after he and family had returned to the US, and stayed with him for another 18 months. This period also was wonderful for me and included: valuable and enjoyable interactions with and learning from him and members of his international group of students and postdocs; more good experiments on Tn5; improvement in my understanding of antibiotics, their actions and resistance mechanisms; and based on Julian’s example, the art of good mentorship. Also, Tn5 was distributed without restriction to laboratories worldwide (patents on such materials were not then in vogue in academia), where it was used by others, for example, to analyze transposition mechanisms, to develop in vitro transposition protocols, and as tools for molecular genetics in other organisms (both prokaryotic and eukaryotic).

In 1977 I moved to a faculty position in Microbiology at Washington University in St. Louis (WUSTL), where my group continued studies of Tn5 and other mobile DNAs for many years, even as my research interests also expanded to studies of Helicobacter pylori and other pathogens. Shortly before my retirement in 2013, I joined with WUSTL colleague Gautam Dantas to help him begin his studies of resistance genes in Latin American urban and rural bacterial populations. My involvement with Gautam also stemmed from the lessons and perspectives Julian had imparted to me many years earlier.

Memories of Patrice Courvalin

After a series of seminars on the east coast my wife Catherine and I landed in Madison at the end of September 1974 and noticed on exiting the plane that the weather was already chilly. This was two weeks after our wedding in France, and it turned out to be the beginning of a three-year honeymoon. I joined the group of Julian, who was on sabbatical in Geneva, and Catherine joined that of Fritz Bach in Immunology.

We appreciated the great hospitality of the Davies family, as, for example, they initiated us into the rites of Thanksgiving and organized numerous parties for the lab members.

Julian was a fine (French) Chef, stating repeatedly that cooking was biochemistry, and that those who were excellent in biochemistry were also excellent in cooking. This, although he invariably had only a yogurt and a banana for lunch. He was deeply Francophile and would never miss a Wales-France rugby game, wishing, of course, that Wales would win. He was much involved in sports, for example as a soccer coach for school children, and traveling between home and lab in Madison almost exclusively by bike, even during a long and terrible winter when he broke a wrist.

Julian was traveling a lot and I remember moving with Catherine to his home during an extended week-end to take care of his children so that his wife Dottie could accompany him. He usually left at the beginning of the week, telling us that he would return on Friday. I never saw him before the next Monday; except for one Friday evening when I was alone in the lab and we had a long scientific discussion.

There were two important devices in the lab. A big Hi-Fi, that for some of us was most unfortunate! Together with Michel de Wilde (from Belgium), we decreed that there be “European time” late in the afternoon, when only classical music would be allowed. This was when most other group members were back home for dinner or, more likely, having beer at the bar across the street.

The second device was an oven, right behind me, with a long list of recipes including the times to prepare one liter of BHI, LB, or MH medium, or agarose, and also to prepare “tarte Tatin pour deux personnes (6 min), quiche Lorraine (4 min), etc.”, and also the lunches I brought in my brown bags. The smell of our cuisine drove my lab mates crazy.

Importantly, Julian was very open to scientific suggestions by his collaborators. For example, he agreed that I start working on antibiotic resistance in the Gram-positive human pathogenic enterococci and staphylococci that I obtained from my previous lab at the Institut Pasteur. Such studies in a group that previously had studied resistance only in Gram-negative bacilli, continued after I left Julian’s lab.

My stay in Julian’s laboratory greatly influenced my career. In addition to my initiation into enzymology and genetic engineering I learned how to run a laboratory and the importance of multidisciplinary collaborations.

Memories of Ólafur S. Andrésson

I was with Julian as a graduate student in biochemistry at UW from 1975 to 1979, and also as a visiting scientist at UBC/TerraGen in 1997. After receiving my Ph.D. I returned to the University of Iceland as a research scientist in the Institute for Experimental Pathology from 1981–2003 and then as professor of genetics in the Department of Biology until my retirement in 2021.

At the end of my undergraduate degree in Iceland, I had studied E. coli ribosomal resistance to antibiotics, and wanted to continue such studies in a good graduate program. I wrote to several leading researchers, and I was immediately hooked on a very welcoming response from Julian in his beautiful handwriting. The UW Biochemistry course program suited me, and there was a good chance of financial support. That is how I joined the Davies lab in the summer 1975, or should I write “the Davies extended family”, with Julian and Dottie caring for each and every member, from undergraduate volunteers and glasswashers (most notably Opal Larson, de facto laboratory manager) to super-postdocs from exotic countries. Social activities acted as a glue; summer picnics, wine and cheese on Friday afternoons (“Goodies”), and Thanksgiving turkey for the stranded foreigners. This was reflected in a smooth working environment with occasional bumps, some chaos and a bit of hoarding. The most notable cultural clashes centered on what music to play in off-hours, and how loud. I sided with the soft classics, folk singers Judy Collins and Joni Mitchell.

Regular lab meetings were fruitful and exciting, something to look forward to, with constructive critique all around, conducted by the masterful hand of the mindful mentor. When it came to writing articles for publication, there was a lot to learn. For Julian it seemed so easy, and no compromise. Both graduate students and postdocs flourished and went on to do well. Julian certainly nurtured a good crop of young scientists in these Madison days.

The whole campus was a hotpot of infectious joy and excitement in exploration and discovery, with an extensive supportive community in the fields of biochemistry and genetics. To top it all, there was the best ice cream in the Midwest at Babcock! These were the pioneering days of recombinant DNA, with the Davies lab contributing restriction enzymes and plasmids, and simultaneously applying hard core biochemistry to studies of protein synthesis and modes of antibiotic action. Maxam-Gilbert sequencing picked up from the Reznikoff lab was particularly enthralling as well as phage lambda cloning, a thread I continued in my Edinburgh postdoc with Noreen and Ken Murray, and then introduced to Iceland.

Midway through my Ph.D., my wife Sigrún and stepson Freyr joined me in Madison. Our little family felt welcome and found caring unselfish support from Julian, lab mates and Reznikoff’s group in the next lab. Summer activities included soccer where Julian was a central organizer. In addition to coaching youngsters, Julian assembled a fierce team recruiting summer visitors from Europe. I once made the mistake of playing against that team, falling victim to a classical English tackle. Revenge finally came when Iceland beat England in the Euro final-sixteen in 2016.

America without a car can be a pain. We found relief in the Davies’s generosity and borrowed the family wagon (not the MG) off and on. And we were ecstatic in the summer of 1979 when we drove the wagon through glorious National Parks all the way to Disneyland, and back through even more wonders. This trip also served Sigrún well professionally in her leading the development of nature interpretation and nature conservation in Iceland.

After leaving the US in November 1979, contact with Julian and Dottie was mainly through Christmas updates. Nearly twenty-two years later, in the summer of 1997, I had the fortune of rejoining Julian for a sabbatical semester when he was a UBC professor and head of TerraGen. So much the same exciting atmosphere as in Madison, leading to new friends and a new research path in lichen genomics. How thankful I am, as are so many others, for the wonderful life of Julian Davies.

Memories of Michel De Wilde

I joined Julian’s lab in February 1976, within days of completing my PhD thesis in the Department of Molecular Biology at the Free University of Brussels. I returned to Belgium in the summer of 1977 after a life changing experience in Madison.

My PhD work under Alex Bollen, himself an alumnus of Julian’s lab, was devoted to the study of bacterial ribosomes, in particular the characterization of mutations leading to antibiotic resistance. When the opportunity to do a post doc in the US occurred, I applied and obtained fellowship support from the European Molecular Biology Organization. The proposed research discussed with Julian prior to my arrival was to extend my studies of bacterial ribosomes to those of yeast. We’ll see how this turned out.

When my wife and I landed in Madison, we were greeted by a smiling and welcoming Julian, who had escaped from the lab’s weekly evening journal club. Next morning offered the first of many cultural shocks: the ground was covered by a foot of snow, an army of snowplows were moving in perfect coordination and hundreds of students were walking toward the campus, each with a backpack. I had not seen anything like that before. The welcome in the lab was warm even if somewhat intimidating. Another cultural shock was the loud playing of the lab’s stereo. Ólafur commented on this.

The question of the exact topic of my research quickly came up and after some procrastination and soul searching, I abandoned the idea of yeast ribosome studies to turn to the lab’s main interest of the lab: drug resistance plasmids! After all, this was the “plasmid era” as Julian wrote in his own autobiography “Gathering No Moss” [Annu Rev Microbiol, 2003; 57; 1–27].

Patrice taught me plasmidology 101; Julian’s graduate student David Smith had isolated restriction enzymes; and so the new tools for studying plasmid structures were in place. Recent findings that plasmids could segregate into “minicells” (small bodies exuded by some E. coli strains) offered new possibilities for studying plasmid gene expression as well. Julian, in his unique way of connecting apparently unrelated dots, put me in contact with Bill McClain and his group in UW’s Department of Bacteriology, where they were studying small RNAs transcribed from T-even bacteriophages during infection. So here we were: I studied the transcription products of the well-studied multidrug resistance plasmid NR1 and pinpointed the location of its coding sequences, although not their functions.

I mention plasmids and restriction enzymes so, yes, I got to clone my first gene while in Julian’s lab. This early experience was to have a great impact on my career.

In February of 1977, Julian had me present my work at a meeting in Jackson Hole, Wyoming. Other than Morimasa Yagisawa putting me on skis there, Marc Van Montagu, a Belgian pioneer of plant molecular biology, liked my presentation and offered me a position in Ghent, Belgium. Under some pressure from family, I accepted. Julian was not too happy about it but did not bear grudges. Indeed, shortly after my return to Belgium, he visited while touring Europe with a group of young soccer players. He also interviewed me for a job at Biogen several years later.

I choose to illustrate some pieces of my stay in Julian’s lab as testimony of his immense qualities and his impact on the careers of all of us co-authoring this obituary.

Memories of Morimasa Yagisawa

I was a visiting researcher in Julian’s laboratory at UW from September 1976 to October 1977. This was soon after the NIH guidelines on recombinant DNA research were released. In accord with them I began cloning the gentamicin resistance gene encoding ANT(2“). I had found that this enzyme adenylylates the hydroxyl group at the 2” position of gentamicin during my Ph.D. research in Tokyo under the direction of Prof. Hamao Umezawa [J Antibiot, 1971:24(12):911–912. https://doi.org/10.7164/antibiotics.24.911]. I was successful in cloning an EcoR1 fragment containing this resistance gene from the large R-factor plasmid pJR66-b into the small multicopy mini-colicin E1 plasmid. This recombinant plasmid pMY26 conferred a level of resistance equivalent to that of the parent R-factor plasmid carrying strain, confirming that the target resistance gene had been cloned.

This recombinant plasmid was later commercialized to serve as a source of ANT(2”) that was used to measure gentamicin concentrations in blood samples. This successful cloning became the first Japanese study to create a recombinant DNA molecule. [J Antibiot, 1979; 32(3):250–253. https://doi.org/10.7164/antibiotics.32.250].

With Julian I also studied the plasmids of antibiotic-producing actinomycetes. I detected two plasmids in the neomycin producer Streptomyces fradiae, pSF1 (14.9 × 106 Daltons) and pSF2 (21.9 × 106 Daltons). Treatment of this strain with acridine dye resulted in derivatives that had lost neomycin productivity, self-tolerance to neomycin and also these plasmids. These facts led to the hypothesis that these plasmids were involved in both neomycin biosynthesis and self-tolerance. [J Antibiot, 1978; 31(8):809–813. https://doi.org/10.7164/antibiotics.31.809].

For me, the success of these two sets of experiments and my mastery of recombinant DNA technology during my short stay with Julian gave me very fond lifelong memories.

Even after I left his lab, Julian continued to provide me with his wise counsel and guidance for many years. For example, he visited my lab in the Tokyo Research Center of Kyowa Hakko Industries Co., Ltd., soon after my return to Japan, and encouraged me and my research group to establish sophisticated systems in industrial biotechnology including recombinant DNA technology.

In 1981, I left the laboratory bench at the Tokyo Research Center and started working as the Managing Editor of The Journal of Antibiotics, an international journal with 34 years history. There also, I was indebted to Julian for his service as a leading editorial board member, and, for more than 25 years, his wise advice to me on the journal’s editorial direction, until my retirement from the journal in 2006. Among many other roles, he served as the Chairman of a study group to give a modern definition of the term “antibiotics” and helped to modernize the definition of antibiotics that had been proposed initially by Dr. Selman Waksman around 1940.

As a senior member of the program committee of ASM’s ICAAC meeting, he also gave me valuable suggestions during the 18 years (1985 to 2002) when I was involved in the conference’s organization. In addition, Julian served as President of the IUMS, and supported the organization of the IUMS meeting in Sapporo in 2011. As I was Secretary-General for the IUMS Sapporo meeting, Julian’s support was extremely encouraging. Patrice Courvalin was also an organizing committee member for these conferences, and this provided nice occasions for renewal of old friendships.

I have a special memory of Julian’s nearly half century of friendship with me. In addition to great discussions of experiments and ideas, our friendship also had important cultural and sportive components. Among them was our slalom together at the Sapporo International Ski Resort during his 2005 visit to deliver a special lecture at the Annual Meeting of the Japanese Society for Bioscience and Biotechnology held in Sapporo. We also had the opportunity to have Julian’s lecture during his 2011 visit at Keio University’s Faculty of Pharmacy, where I was a professor in fundamental science.

Memories of Thomas J. White

I received my PhD in 1975, studying mechanisms of molecular evolution at the University of California in Berkeley. In my first as a postdoc at the University of California San Francisco Medical Center I investigated the role of environmental mutagens in breast cancer. During this year I met my future wife Leslie, who was quite unhappy that I was planning to leave Berkeley for 3 years in Wisconsin. Nevertheless, we arrived in Madison in mid-May 1977; Lake Mendota still had some floating ice, but the weather shortly became hot and humid – as if “Spring” was limited to 2 weeks. Leslie soon returned to California to teach in San Francisco and we maintained our relationship via multiple trips across the country by car, plane and train.

Julian’s lab was a mélange of graduate students and postdocs from multiple countries. The ATCC sent me a culture of “E. coli” that didn’t look like the one I knew. Patrice Courvalin taught me how to recognize Pseudomonas aeruginosa by its cadaverous smell, and not to trust the ATCC without confirming it myself. Based on my background with molecular phylogenetic analyses, Julian and I settled on experiments to test his theory of the origin of antibiotic resistance genes. If the protein sequences or immunological relatedness (remember, this was before easy DNA sequencing) of the enzyme from the producing microbe was closer to that from a resistant enterobacterial species than were other proteins from those two species, it would support his theory of horizontal transfer. If differences were similar to those expected for descent from a common ancestor, this would indicate that his hypothesis was likely incorrect. Some initial immunological comparisons showed the Streptomyces enzyme did not cross-react with antisera against an aminoglycoside enzyme isolated from a resistant E. coli, but surprisingly, the homologous enzyme from a Gram-positive bacterium did cross-react. This was an initial hint that Gram-negative – Gram-positive gene transfers could occur – an interpretation that Patrice and group showed directly in lab experiments back at the Institut Pasteur.

Julian and Dottie made Leslie and me very welcome in Madison. I also recall wonderful Japanese dinners prepared by Morimasa and his wife Mieko. Julian taught me to play squash, and invariably beat me with his 3 wall return shots. But after a year away from Berkeley and Leslie, I began to apply for academic positions in the Bay Area. No biochemistry openings were available at UC Davis, UC Berkeley, UCSF, etc, but the biotech companies Cetus and Genentech were hiring. I joined Cetus in July 1978 and eventually became VP of Research. After the Polymerase Chain Reaction was invented by Kary Mullis, I left Cetus in 1989 to join Hoffman-La Roche’s pharma group in Nutley, NJ and subsequently became Senior VP of R&D of Roche Molecular Systems and then Chief Scientific Officer at Celera Diagnostics.

Over the almost 50 years since my time in Julian’s lab, I met with him a few times at UBC and ASM conferences. At one dinner in San Diego with members from this article, I distributed cheek swabs for everyone to submit a saliva sample for genetic ancestry profiling. Julian was definitely of Welsh ancestry.

During the 1990’s I had the great pleasure of becoming friends with Morimasa’s son Tomomasa, and Patrice’s sons Tom and Damien while they were doing graduate studies at UC Berkeley.

Family life recounted by Vicky, Robin and Jeremy Davies

Our father was a big presence throughout our lives. When we were young, he enthralled us with stories of his childhood. Our imaginations would run wild hearing about the time he spent in Wales during WW II looking for shrapnel on his bike and selling stale ice cream cones to his chums when sweets were scarce. It told of his resilience and resourcefulness that got passed on to us.

Our first taste of the good life was in Paris in the 1960s: French bakeries, crèpes, better wine for the adults, castles, art museums, and beautiful parks (though we children found it difficult to stay off the grass). The sabbatical he took in 1974 in Geneva Switzerland exposed us to skiing in the Alps, cheese fondue, and the international nature of the city. Our French improved, he was envious of our ability to adapt so quickly. No surprise that Vicky found her calling in the culinary world.

Good health was impressed upon us and that included participation in sports. He rode his 3-speed bike to work all season long, played competitive squash, and got us to the ski slopes whenever possible. To this day, we all ride our bikes, enjoy the mountains for camping, hiking or skiing, and Robin & Jeremy still play hockey.

He always included others and asked for our participation. There was a constant influx of department visitors and students to dinners at our home. When Julian’s father Norman came to live with us his presence added a depth to the household dinner discussions and the smell of Norman’s pipe something we can never forget.

Julian worked and traveled a lot, yet somehow found time to bring each of us a small gift from his international meetings. He bought a red MG that Vicky & Robin got to drive to High School; this was a vast improvement over the orange station wagon referenced by Ólafur Andrésson. By far the biggest gift was his encouragement and support in our lives. The rich experiences the entire family has shared, whether happy or sorrowful, gave us deeper connection and a sense of inner strength.

As his granddaughter Madeleine wrote in his obituary: “While Julian possessed the sharpest of wits, he was also capable of immense feeling. He expected and hoped for those around him to experience their fullest potentials and would do what he could to help them reach it.” The family remembers him as a passionate working man and a kind family man. His lab and scientific community gave him so much. He always pushed for a little bit more from each of us while accepting who we were and who we became. His joyous nature will remain in our hearts.