Copyright � 2009 by the Genetics Society of AmericaDOI: 10.1534/genetics.109.101360

Perspectives

Anecdotal, Historical and Critical Commentaries on Genetics

Teach, Then TrustElizabeth W. Jones (1939–2008): Mentor to Many

Tracey DePellegrin Connelly, Sandra K. Lemmon,* Aaron P. Mitchell† and John Woolford, Jr.1,†

*Department of Molecular and Cellular Pharmacology, University of Miami, Miami, Florida 33101 and †Department of Biological Sciences,Carnegie Mellon University, Pittsburgh, Pennsylvania 15213

FOUNDATIONS

Elizabeth (Beth) Jones reveled in the excitement ofscientific discovery and fostered curiosity with stead-fast determination. With high standards and rigor, sheapplied a firm but gentle hand to cultivate the curiosityof generations of students. Inquisitiveness aboutscience and the workings of nature was threadedthrough her life, unabated, from early childhood.

Born in Seattle, Beth spent her first decade in Diablo,a small town at the base of the Washington Cascades, inthe Skagit River Gorge, where her father worked forSeattle City Light at the Diablo Dam Powerhouse.Lacking even a highway—trains provided the soleaccess—the setting provided Beth and her three siblingsfertile fields for adventure. Educated in a one-roomschoolhouse (Figure 1), they spent their days playingalongside the dam, building, climbing, and exploringthe environs of Diablo Lake.

Beth’s parents set her stage, encouraging reading,goal setting, and risk taking. Their tenet—teaching andthen trusting—came to exemplify Beth’s philosophy asan educator, an editor, and an experimentalist.

In an autobiographical lecture presented at CarnegieMellon University last March, Beth acknowledged the

remarkable influence her parents had on her. Her mother,Dorothea (‘‘Dowty’’), was intrigued by plants and flowers;her father, ‘‘KC,’’ was an avid bird-watcher. The familytraveled extensively throughout the West: Yellowstone, theGrand Canyon, Mesa Verde, Crater Lake. Beth said herparents ‘‘provided a background of heightened aware-ness of things around me that fed my curiosity.’’

Her father felt his children should reach for whateverthey wanted, regardless of their gender. His influencecan be seen throughout Beth’s life, especially in the wayshe mentored women scientists.

Beth’s early schooling sowed the seeds of her teachingphilosophy. She learned to read at an unusually youngage and confessed that she ‘‘became very cocky about[reading].’’

Beth admired her teacher in the one-room school-house, because she managed to keep track of theprogress of every student and accommodate each one.Mrs. Kerchen ‘‘lay in wait’’ as Beth later described it. Sheonce asked Beth to read aloud a selection containingthe word ‘‘aisle,’’ which Beth had never before seen andhad no idea how to pronounce. She stumbled in thereading and thereby learned a lesson about beingoverconfident. As Beth’s students (and many of her

It was a very dangerous life. We were always out fishing,playing in the riverbed. My parents had to teach us andtrust us.

E.W.J.

Even if the open windows of science at first make us shiverafter the cozy indoor warmth of traditional humanizingmyths, in the end the fresh air brings vigor, and the greatspaces have a splendor of their own.

Bertrand Russell, What I Believe, 1925

My father, in a mental sense, raised me as a son. Althoughhe had southern roots, he did not subscribe to stereotyp-ical ‘boy things’ or ‘girl things’; rather he expected me todo well in whatever I did and to seek out and make themost of opportunities . . . I didn’t realize for a long timewhat a gift that was. I decided what I wanted to do basedupon what I wanted to do . . . and didn’t decide based onwhether it was proper.

E.W.J.

1Corresponding author: Carnegie Mellon University, Department ofBiological Sciences, Mellon Institute, 4400 Fifth Ave., Pittsburgh, PA15213. E-mail: jw17@andrew.cmu.edu

Genetics 181: 357–365 (February 2009)

colleagues) will recognize, she adopted some of Mrs.Kerchen’s teaching style.

Diablo had no high school, so the family relocated toa larger town when Beth’s elder sister reached eighthgrade. Middle school in Longview, Washington, heldfew challenges for Beth, who was nearly 2 years aheadin her lessons. She challenged herself by learning toplay (and quickly mastering) the French horn. She wasone of the 5 (of 162) students in her high schoolgraduation class who went off to college. (One of themwas John Abelson, Cal Tech Emeritus Professor andnow on the faculty of the University of California, SanFrancisco. Abelson recalled Beth setting an extremelyhigh standard of performance in physics and chemistrycourses.)

TRANSITIONING TO A CAREER IN GENETICS

Beth had aspirations to be an organic chemist andchose chemistry for her major at the University ofWashington. But her interest in that subject soonwaned, and she increasingly focused on zoology, despiteher distaste for taxonomy.

Beth soon became aware of a glass ceiling in science.Despite her having just earned the highest grade on thefinal exam in his accelerated chemistry course, one of

her professors told her that there was no room for awoman in chemistry. Beth agreed with him and began tolook elsewhere for a career.

She became interested in genetics while washingglassware in Herschel Roman’s yeast genetics lab andenrolled in every genetics course she could find. Sheemerged from the University of Washington in 1960with a bachelor’s degree in chemistry, looking tobecome a geneticist.

Roman picked Beth to be one of the first students inthe new Genetics Ph.D. program he established in 1959at the University of Washington. Beth said Roman—whom she described as a ‘‘goader, who always kept youmoving with his tongue’’—was the most influentialmentor in her life. She credited Roman and his wife,

Figure 1.—Beth’s one-roomelementary school, Diablo, Wash-ington.

I never really thought much about what I wanted to do inthe long term; I always lived in the moment. In college, Iinitially had no idea there was such a thing as graduateschool. Herschel Roman and his wife, Caryl, began toopen my eyes, and I learned more about graduate schooland what it was about. I entered the genetics Ph.D.program at the University of Washington, but even thenI didn’t think about what I wanted to do with it. Continuingmy education might have in part been the deferral of adecision. One of the few legitimate career options forwomen at the time was teaching, and I was being pushedpretty hard in that direction by my parents, my teachers,and by everyone I encountered. My decision to be achemist and then a geneticist was perhaps one way ofresisting having my decisions made for me.

E.W.J.

That’s the kind of teacher she was—very encouraging, butshe didn’t let you get by anything.

E.W.J.

358 T. DePellegrin Connelly et al.

Caryl, with broadening her perspectives outside ofscience and introducing her to literature. Caryl recom-mended novels for Beth to read, starting with MaryRenault’s books about ancient Greece. Beth was knownfor immersing herself into a broad expanse of non-fiction subjects and novels, notably mysteries; her bookcollection numbered well into the thousands.

Beth’s graduate work centered on the question ofwhy mutations in different ADE genes arose at differ-ent frequencies. Beth sought to test the simple hypoth-esis that mutant frequency reflected relative gene size.But how does one measure gene size? Beth initiallyaddressed the question by generating intragenic mei-otic maps, for which the data were workable, though‘‘not very pretty’’ ( Jones 1972). Later she generated X-ray mapping data that were in agreement. It was clearthat gene size could not account for the differences inmutant recovery rates.

In 1964 Beth received one of the first doctoratedegrees awarded by the storied department thatplayed a major role in making yeast genetics main-stream science. She emerged a skilled practitioner ofgenetics.

DEVELOPING A TEACHING PHILOSOPHY

Beth moved on to postdoctoral training at MIT withBoris Magasanik, who fueled her interest in under-graduate teaching and research. It was there that Bethfirst learned to ‘‘do science in a more complicatedfashion.’’

MIT provided the setting for Beth’s pioneering work indeveloping standards for a challenging, stimulating,and engaging undergraduate research experience.After 1 week teaching a recitation session for under-graduates as part of Maury Fox’s genetics course, Bethrealized she was covering the same ground as Fox,despite their previous arrangement that she wouldexpand on topics he wouldn’t discuss in class. Thestudents were not engaged; neither was Beth.

So she decided to ‘‘make it up as (she) went along’’ andintroduced the students to the current scientific litera-ture, settling on the genetics of bacterial viruses, to greatsuccess. People from other discussion sections driftedinto her recitation. Student feedback was enthusiastic.‘‘Reading original literature was wonderful for the

Figure 2.—Beth and co-instructor Peter Berget, sur-rounded by graduating seniorswho had joined her for the2003 Summer Research Institute.Reprinted with permission fromCarnegie Mellon University.

I hadn’t expected to go [to MIT]; opportunities forwomen at that level were less available because of theexpectation that they would quit. I got the chance becauseJon Gallant, a junior faculty member in genetics at theUniversity of Washington, insisted, under interrogationby my postdoc mentor, that I would stick it out.

E.W.J.

I didn’t know it was fashionable at the time for postdocs tohave nothing to do with teaching. In fact, if you’re goingto do it, you should ask your mentor. But I just accepted it[when Maurice had asked me to teach], and did it.

E.W.J.

Perspectives 359

students. They were getting the real deal,’’ Beth said.‘‘They felt like grownup scientists.’’

Beth’s other teaching experience at MIT spawned thebeginnings of a fruitful endeavor, one that wouldcontribute to changing laboratory biology courses andthe undergraduate science experience. The MITadmin-istration and biology department had been dissatisfiedwith the current structure of the cookbook, formulaiclaboratory courses. Spurred by Magasanik, David Bot-stein was promoted to faculty status (nontenure track)and asked to redesign the course. He wisely chose Bethto help him. ‘‘I wanted to do phage, Beth wanted to doyeast,’’ Botstein, now Professor of Genomics at Prince-ton University, said. ‘‘We compromised on bacteria.’’

The two young scientists labored over the coursedevelopment. What would be the most effective way toengage students in the excitement of science andinspire them? David and Beth decided to have studentsperform real science—experiments without a predeter-mined outcome or guarantee of success. The instructorsbrought a project to students and offered protocols asstarting points. The students designed, carried out, andanalyzed their own experiments. The first course—withopen labs running 14 hours each day, 7 days a week—was a resounding success. Captivated by actually doingreal science, the students invested endless hours to learnan unknown outcome.

David and Beth also wanted the students to experi-ence a real scientific outcome: a story published in apeer-reviewed journal. Indeed, students from that firstproject lab published their discoveries (Botstein andJones 1969). Botstein recalled that the Journal ofBacteriology refused to include the undergraduates’names as coauthors. (In contrast, as Editor-in-Chief ofthis journal, Beth strongly supported inclusion of un-

dergraduate coauthors.) But the instructors successfully‘‘cajoled and negotiated’’ and the names of 24 un-dergraduate students appeared in a footnote on thearticle’s first page. Beth and David invented the firstproject lab at MIT, which to this day is the model forbiology labs at MIT, Carnegie Mellon, Princeton, andother universities around the nation.

Beth continued to provide these experiences to un-dergraduates throughout her career at Carnegie MellonUniversity. As one of the first Howard Hughes MedicalInstitute (HHMI) Professors, she was awarded over $9million to apply research-grade thinking to teachingand to create novel educational programs.

She developed the Summer Research Institute, whicheach year immersed 12 promising sophomores in re-search (Figure 2). In 2007 she and her studentspublished an article reporting the novel finding thatyeast mutants resistant to the amino acid analog 5-fluoroanthranilate acid carry mutations in any of thefour TRP genes required for conversion of anthranilateto tryptophan (Jones et al. 2007). Beth saw this articleand the one published nearly 40 years earlier, with 66undergraduate coauthors in between, as her bookendundergraduate-authored publications.

HER HEART WAS IN THE WORK:AT CARNEGIE MELLON

Beth joined the faculty as an Assistant Professor ofBiology at Case Western Reserve University in 1969. Shelater candidly acknowledged that the department wasnot the right place for her, setting herself a deadline‘‘either to leave the place or to get out of science.’’

In 1974 she joined the Department of BiologicalSciences at Carnegie Mellon University, becoming thefirst female faculty member in the department. She

Figure 3.—Beth in her officeat Carnegie Mellon. Photo is byKen Andreyo. Reprinted with per-mission from Carnegie MellonUniversity.

360 T. DePellegrin Connelly et al.

immediately fell in love with the University, never losingthat sentiment (Figure 3).

During this time, issues of gender inequality hadcome to the fore. Beth considered herself first andforemost a scientist. But among other scientists, sheacknowledged the limitations and challenges that beinga woman in an all-male field brought. With character-istic aplomb Beth faced the prospect that she had beenhired to fill a quota for X chromosomes.

During an autobiographical lecture in 2008, Beth toldthe audience that when she was a young scientist she andother women scientists had fewer choices—which,candidly, meant a dichotomy of family or career. Sincethen, she said, times have changed for women in science.But Beth lived life on her own terms and was adamantthat her choices were in fact just those—her own.

Beth and the late Bill Brown set about establishing anexciting curriculum. The two-semester lab course forundergraduate juniors that Beth established has re-

mained the foundation of the curriculum. Beth de-signed and taught the department’s Advanced Geneticscourse. And she excelled at it. Beth brought sophisti-cated discussions of complex genetics to undergradu-ates and grounded them in the principles so they couldunderstand how the investigators were thinking andhow their thinking evolved—how they revised theirhypotheses and made discoveries. She taught herstudents to think like geneticists. Her colleagues re-alized how important teaching was by watching Bethdo it.

Beth is remembered by many for her enthusiasm forteaching and turning young people on to the processand the excitement of discovery, and for providing amilieu in which each could thrive. She had an in-dividual, personalized manner that resonated withstudents. She memorized photographs of her students(by the second day of class!) so no one could hide underan anonymous cloak. She had a talent for figuringout ways to draw even the quietest student into theconversation.

She served in something of a maternal role for manystudents, acting as a conduit between them and facultymembers. Late night parties at Beth’s house wereterrific fun and happened often. She always had a fulllineup of board and parlor games to play. She activelysought out games that might pique the interest (andreveal the skills) of some of the lab wallflowers. The

Figure 4.—–Beth engaged with studentsin the lab. Reprinted with permission fromCarnegie Mellon University.

Most of the time I couldn’t assess what I had earned onmerit, and what I had earned because they needed a body. . . The way I coped with that—so they chose me for thewrong reason, [but] they got lucky and they got thecompetence they should have been looking for.

E.W.J.

Perspectives 361

parties often went on until 2 am or later; Beth was usuallythe last one standing.

Inspired by her teaching experiences at MIT, Bethbrought undergraduates into her lab, involving them inher research. She continued to do that throughout hercareer, inspiring others to follow her lead, firmlyestablishing the practice in the department. More than130 students passed through the doors of her lab;several of them, like Roy Parker, Professor at theUniversity of Arizona, rose to the highest levels ofacademic science.

Beth encouraged students to take the initiative indoing research. Undergraduate researchers in her labwere expected to earn their place. Their initiation wasmedia and reagent preparation; if their curiosity waspiqued by what was happening around them, they wereassigned a graduate student or postdoc mentor. Bethmentored the mentors, paying forward her passion forresearch. The students and postdocs she trained are herlegacy (Figure 4).

In the classroom and in the lab, Beth demandedcompetency, originality, and unwavering ethics. Herstandards were driven by her respect and integrity forthe purity and intellectual honesty of the science. Herlegendary toughness was leavened with a brusque senseof humor, and with patience, and with encouragement,particularly of students she sensed were agile, inter-ested, and engaged. It was always a joy to come to Bethwith new and interesting results, because one coulddepend on her to share in that excitement. She thrivedon the freedom offered by exploration just outside themargins or on a not necessarily linear path to un-derstanding. She lamented when students (or col-leagues) took a safer route rather than extendthemselves beyond their zones of comfort.

As director of the HHMI Undergraduate BiologicalScience Education Program at Carnegie Mellon, Bethestablished numerous innovative educational pro-grams, coalescing in elegant fashion the roles ofscientist and teacher. To fully understand genetics,students must think at a higher cognitive level. Thistype of mastery requires innovative problem-solvingapproaches, in addition to practice that reinforcesscientific concepts and processes. Along with CarnegieMellon Biological Sciences faculty Linda Kauffman andCarnegie Mellon Senior Research Scientists Albert Cor-bett and Benjamin MacLaren, Beth developed the Ge-

netics Cognitive Tutor, interactive software designed togive students a boost in problem solving and thinking.

Beth’s balanced approach to mentoring fostered in-dependence in a nurturing environment. She was awonderful mentor who took a long view of her students.Rather than micromanaging her lab, she allowed—in fact,expected—independent thinking. She gave the youngerscientists a framework for designing experiments, buteveryone eventually worked on his or her own. Roy Parkersaid ‘‘She had a huge heart and was all about teaching thestudents about science, but with intellectual honesty,integrity, and devotion. There were no shortcuts.’’

TO DISCOVER: BRINGING GENETICSTO CELL BIOLOGY

At Carnegie Mellon Beth developed a new area ofresearch on protein degradation. This led her topioneering work on the structure and function of theyeast lysosome-like vacuole and on membrane traffick-ing. She developed overlay assays for protease activitiesand often expressed her delight at what an interestingstew one could cook up on a plate of colonies. Shecredited the Ames test, in which liver extract is overlaidon a petri plate, as a ‘‘liberating’’ influence.

Her genetic screens for proteinase-deficient pep mu-tants were surprising because they defined many moregenes than necessary to encode a structural gene or oneor two regulatory genes. Many exhibited pleiotropicphenotypes, including defects in multiple proteaseactivities, amino acid analog hypersensitivity, and spor-ulation deficiency. Her seminal article on discovery ofthe pep mutants, published in Genetics ( Jones 1977),appeared before the vacuolar localization of theseenzymes was known. Beth speculated that these pleio-tropic mutations might ‘‘cause changes in the structureof the compartments containing these enzymes’’ or‘‘might alter components of the system that places theenzymes in the compartments’’. Indeed, the idea thatPEP gene products might govern protease compartmen-tation was an insight that proved prescient. She hadidentified the first genes encoding components of thevacuolar protein sorting machinery.

Among Beth’s pep mutants were ones unable to sendvacuolar hydrolases to the proper compartment, onesunable to acidify vacuoles, and ones unable to formdetectable vacuoles. Isolation of the PEP genes andcharacterization of the products by Beth and othersuncovered many of the components of vesicle traffick-ing, including syntaxins, Sec1/Munc18 family proteins,and Rab effectors, as well as other vesicle componentsnecessary for fusion at different stages of the vacuolarprotein sorting pathway (e.g., retromer, CORVET, andHOPS complex subunits).

Complicated genetic challenges energized Beth. Thistrait enabled her to discover and explain phenotypic lag

What’s far more important is that [those in the lab] bringin their experimental problems—things they can’t get towork or can’t figure out—and as a group we try to figureout where the problem is, how it can be overcome or whatit may mean. The interplay of many minds often leads usto places we couldn’t get to by ourselves.

E.W.J.

362 T. DePellegrin Connelly et al.

of the vacuole, in which a mutation affecting a singlehydrolase might not reveal itself for many generationsuntil the hydrolase and its targets became depleted.This mysterious ‘‘epigenetic’’ phenomenon led to theuncovering of the basis of proenzyme processingcascades and autocatalyic protease activation withinthe vacuole and lysosome.

Her lab also studied aspects of metabolic regulationof the vacuole, particularly the upregulation of vacuoleconstituents during nutrient deprivation. Her findingthat pep mutants cannot sporulate provided hints of howcellular remodeling and recycling of cellular contentsare affected by lysosomal dysfunction. Today we knowthat the lysosome/vacuole is critical for cells duringstarvation and autophagy.

Beth’s research on the vacuole began during anexciting period in the development of yeast as anexperimental organism, and she was one of the firstpeople to bring genetic analysis to bear on issues of cellbiology. She continued to work on the PEP genes andvacuolar function for 30 years. Her observations ofyeasts were seminal and had a major impact on ourunderstanding of endosome/lysosomal function andgenetic disorders of these organelles.

TO SERVE: DEPARTMENT HEAD,WRITER, EDITOR, REVIEWER

As department head, Beth was a strong advocate forher faculty, defending their goals and interests andlooking out for their welfare. She was well known forapproaching Deans and Provosts and Presidents with awell-thought-out plan, which she would defend logicallyand emphatically. She tirelessly pursued resources todevelop her department, engaged faculty in decisions,led efforts to bring computational biology and neuro-biology into the department, and strengthened cell anddevelopmental biology. Beth demonstrated diplomacyas department head and gathered consensus, even asshe voiced her opinion.

Beth remained steadfast in her core belief thatinformation must be presented clearly and accurately,regardless of framework or purpose—written or oral,formal or informal. She had a well-deserved reputationas a stickler for grammar. Beth viewed correcting flawedwriting almost as a moral calling. She sent departmentale-mails pointing out recurring stylistic and grammaticalerrors—one memorable evening message declared that‘‘the misuse of the phrase ‘begs the question’ is rampantin our department.’’ Precision and pith were prized—no doubt contributing to the passion and success of herendeavors as writer and editor.

Beth’s service to the genetics community was carriedout with her usual depth of commitment. From 1990 to1993, she served as Chair of the National Institutes ofHealth Genetics Study Section (she participated formany years prior), for which she read every proposal

and actively participated in each review. She served as anAssociate Editor of the Annual Review of Genetics forover 15 years. And she served as Associate Editor onnumerous editorial boards, including Genetics (for 20years), Yeast, and Molecular Biology of the Cell. WithJim Broach, John Pringle, and Jeff Strathern, Bethedited ‘‘The Yeast Books’’ in the Cold Spring HarborMonograph Series ( Jones et al. 1992).

Beth coauthored with Dan Hartl Genetics: Analysis ofGenes and Genomes, now in its seventh edition publishedin 2008 (Hartl and Jones 2008b). They also partneredto author Essential Genetics: A Genomic Perspective (Hartl

and Jones 2008a). Both textbooks continue to provide asolid introduction to modern genetics without beingoblivious to historical context.

‘‘I was always impressed with the precision andexactitude of thought that she brought to teachingand writing about genetics,’’ said Hartl. ‘‘Beth realizedthat progressively dumbing down the curriculum didnot make the subject more accessible to unpreparedstudents, but merely bores and alienates those who wereprepared.’’

Beth’s numerous distinctions included being the firstwoman at Carnegie Mellon to be appointed the Freder-ick A. Schwertz Distinguished Professor of Life Sciences.She also received the university’s Robert Doherty Prizefor Excellence in Education and the Julius AshkinTeaching Award. In 2008, she received the inauguralExcellence in Education Award from the GeneticsSociety of America and also its Lifetime AchievementAward (Wright 2007) for her pioneering work in yeastgenetics.

Beth’s values in education, language, genetics, andservice to the scientific community dovetailed duringher 12 years as Editor-in-Chief of the journal Genetics,from 1996 until her untimely death. Under her steward-ship, the journal grew—literally—in size, scope, and itseditorial board. With a belief in an inclusive approach topublishing science, Beth encouraged submissions ofarticles on myriad topics and brought internationalrepresentation to the editorial board. Long held as themost prestigious venue for population and evolutionarygenetics articles, submissions to Genetics doubled inall areas during Beth’s tenure as Editor-in-Chief. Shemade sure that the journal maintained the higheststandards in the midst of its burgeoning growth. Beth’sreputation for integrity and fairness was well knownamong her team of associate editors, whose numberspeaked at nearly 90.

Beth pointed to the constant growth of the journal asevidence of the health of and interest in model organismgenetics. When the journal published especially largeissues, usually in December and sometimes topping 60articles, she would hoist the printed journal into the airand remark on the stamina and dedication of her editors;she referred to the busiest associate editors (those han-dling an exceptional number of manuscripts; you know

Perspectives 363

who you are . . .) as her ‘‘heroes.’’ Above all, Beth was mostproud of the quality of the articles.

Beth believed in solid science, and whether an articlewas cited 1 year or 10 years after publication, she sawvalue in the journal’s role of vetting and tellingsignificant stories and as a repository of discoveries thatcatalyze further advances. Beth saw to it that the entirejournal content (dating to the journal’s origination in1916) was placed online in a searchable, free archive—acostly but worthwhile and well-used resource. In theincessant debate over the importance of impact factor vs.the intrinsic value of an article, she would often counterwith the example of Barbara McClintock.2 Beth had littletolerance for the popular-but-fleeting or ersatz articles.

Beth modernized the journal in several critical ways.In 1996 when she succeeded retiring Editor-in-Chief JanDrake, peer review was accomplished using hard copiesof manuscripts passed around by the United StatesPostal Service, many of them languishing too long intransit. The journal, too, was a print-only effort. Bethand then-Managing Editor Leah Kauffman transitionedthe journal into the digital age, first with electronicrecord keeping and then partnering with HighWirePress to provide an online edition of Genetics. AndBeth was an early proponent of open access to scholarlyresearch, making an online version of each articleavailable within 3 months of its publication and estab-lishing the journal’s publish-ahead-of-print policy in2004.

Beth took great pleasure each year in choosing thecolors of the journal cover. Kauffman recalled Beth’sfirst choice of journal cover: a bright shade of pink. Inlater years, she pored over hundreds of Pantoneswatches, asking colleagues for opinions and analyzingthe implications of each color. In 2006 when thejournal’s Board of Senior Editors, led by SuzanneSandmeyer, transformed the journal’s visage, Bethrelished the opportunity to present a fresh face thatshowcased the science underneath.

Ever the wordsmith, Beth’s attention to detail knewno bounds. She poured over nearly every acceptedmanuscript prior to its publication. Some of her editorswondered why she was compelled to read more than

7000 manuscript pages each year, knowing her respon-sibilities as Editor-in-Chief and Chair of her Departmentand her HHMI work, her teaching, and more. She saidsimply, ‘‘I want to see everything that we publish.’’

And see she did. Beth’s vision entailed tremendousresponsibility—to the authors, primarily, and to hereditors, readers, reviewers, and the scientific commu-nity. She felt it important to review most manuscriptssubmitted to the journal, to give each one a fair shake.She made strides in improving time in review and timeto publication. She balanced a commitment to publish-ing sound science while allowing the largest possibleaudience the easiest ways to access the articles. Bethknew what she knew and sought input from editors andcolleagues in cases in which she lacked expertise. Evenwhen she held strong opinions about a topic, shedemonstrated a capacity to render objective decisions.And she apologized when she erred.

No matter how esteemed a journal, scientific publish-ing is an environment in which the sacred and theprofane, the intellectual and the practical, collideduring weeks filled with unrelenting deadlines, scathingcomplaints, and disgruntled authors. Whatever thecause, the solution for Beth involved very nearly thesame pattern: a quick but thoughtful and thoroughreview of the situation, followed by a succinct presenta-tion of the solution. Beth’s command of the language,coupled with her high character and enduring strength,gave rise to e-mails and phone calls that remainlegendary, if only to the recipients. Consider her e-mailto an author who had omitted necessary clarifications atproof stage. ‘‘There will be corrigenda,’’ she wrote.‘‘They will be honest. We can discuss the wording.’’

LEAVING A LEGACY

‘‘The most important thing in science,’’ Beth said onmore than one occasion, ‘‘is the people.’’ Beth lived bythat aphorism. She spent her life cultivating studentsand cultivating the network of relationships that create ascientific community. The spark of individual curiositywas the essential unit of discovery, and a web of mentorsand colleagues provided the oxygen to fan it into flame.

Echoes of Beth’s hearty laugh—and her trademarksigh when irritated or impatient—reverberate with hermany students, colleagues, and friends. Her legacy ofreinvigorated, dynamic science education already livesbeyond her own students and even her own field, in thegenerations learning science not as a body of knowledgeto absorb, but as a discipline of thought and discovery,consummated through collegiality.

Notes: Excerpts from this piece were taken fromseveral sources: The Elizabeth W. Jones MemorialSymposium held on October 15, 2008, at CarnegieMellon University; Beth’s taped presentation ‘‘Making itUp as I (We) Went Along’’ from The Journeys Lecture

2Referring to McClintock’s work in the genetic regulation of the lacoperon, which McClintock had demonstrated in 1951 but that did notcome to light until the 1960s.

If you believe [the manuscript] to be unsuitable forGenetics and you have the review to back you up, goahead and reject it. You as [associate editor] are not asecretary to reviewers. When I was an AE I even rejectedmss that both reviewers said should be accepted because Ifelt that the reviewers had missed a key element. So gowith your best judgment. I will back you up.

E.W.J, in a note to a Genetics Associate Editor

364 T. DePellegrin Connelly et al.

Series at Carnegie Mellon University on March 31, 2008;Beth’s own words from her first-person profile inJourneys of Women in Science and Engineering: No UniversalConstraints (Ambrose et al. 1997); and personal corre-spondence and electronic mail from Beth to the authorsand others.

We sincerely thank Kristin Boise, David Botstein, Amy Burkert,Shelley Esposito, Stanley Gartler, Leah Kauffman, Jon Jarvik, MaryAnne Jarvik, David Jones, Jennifer Sciullo, and V. Emily Stark for theirinsights, clarifications, and contributions, and others too numerousto mention for providing stories and memories of Beth. We appreciatethe use of original photographs owned by Carnegie Mellon Universityand by David Jones.

LITERATURE CITED

Ambrose, S. A., K. L. Dunkle, B. B. Lazarus, I. Nair and D. A.Harkus (Editors), 1997 Journeys of Women in Science and Engi-neering: No Universal Constraints. Temple University Press,Philadelphia.

Botstein, D., and E. W. Jones, 1969 Non-random mutagenesis ofthe Escherichia coli genome by nitrosoguanidine. J. Bacteriol.98: 847–848.

Hartl, D. L., and E. W. Jones, 2008a Essential Genetics: A GenomicsPerspective, Ed. 7. Jones & Bartlett, Boston.

Hartl, D. L., and E. W. Jones, 2008b Genetics: Analysis of Genes andGenomes, Ed. 7. Jones & Bartlett, Boston.

Jones, E. W., 1972 Fine structure analysis of the ade3 locus in Saccha-romyces cerevisiae. Genetics 70: 233–250.

Jones, E. W., 1977 Proteinase mutants of Saccharomyces cerevisiae.Genetics. 85: 23–33.

Jones, E. W., J. Pringle and J. R. Broach, 1992 The Molecular Biologyof the Yeast Saccharomyces, edited by J. N. Strathern. Cold SpringHarbor Laboratory Press, Cold Spring Harbor, NY.

Jones, E. W., P. B. Berget, J. M. Burnette, III, C. Anderson, D. Asafu-Adjei et al., 2007 The spectrum of Trp� mutants isolated as5-fluoroanthranilate resistant clones in Saccharomyces bayanus, Sac-charomyces mikatae and Saccharomyces paradoxus. Yeast 25: 41–46.

Wright, R., 2007 The 2007 Genetics Society of America Award forExcellence in Education. Genetics. 175: 467–468.

Communicating editor: M. Johnston

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