Kiki Milner as Viola in the PAD’s production of Twelfth Night, or What You Will April 17-20. Photos by Whitney Curtis/WUSTL Photo. Hires versions upon request.

Spoken from a modern proscenium, or ‘picture frame,” stage, the soliloquy feels hushed and contemplative, a quiet window into private thoughts.

But William Shakespeare did not write for the proscenium. His Globe Theatre had no curtains or spotlights. Groundlings crowded at the actors’ feet, sharing the sun, elbows propped upon the platform.

“When Hamlet or Macbeth asks what he should do, it’s not a rhetorical question,” says Kate Drummond, who recently defended her senior thesis on “Now I am Alone: A study of Isolation in Soliloquies.” “He wants an answer.”

This month, Drummond and the Performing Arts Department in Arts & Sciences will celebrate the 450th anniversary of Shakespeare’s birth with a new production of Twelfth Night, or What You Will. Though best known for its playful satirizing of gender roles, the beloved comedy also offers a window into Shakespeare’s development of the soliloquy form.

“Shakespeare likely wrote Twelfth Night in the same year he wrote Hamlet,” says director Henry Schvey, professor of drama. “Viola has a number of soliloquies, but there’s also the unique phenomenon of Malvolio’s soliloquy, which is overhead, and commented upon, by other characters.”

In other words, “Shakespeare is riffing,” Schvey says. “He’s flexing his artistic muscles. He’s already questioning the convention he helped establish.”

Ariel Saul as the singer.

First performed in 1602, Twelfth Night centers on Viola, a shipwrecked traveler grieving for her lost twin, Sebastian. Alone in a strange country, Viola adopts male attire and a new name — “Cesario” — and enlists as a page to noble Duke Orsino, with whom she soon falls in love.

But Orsino pines for the Countess Olivia, who has sworn seven years’ chastity while mourning her own lost brother. At wits end, the Duke dispatches Cesario to woo on his behalf and Olivia does fall in love — with Cesario, much to Viola’s chagrin.

Matters are further complicated by Sir Toby Belch, Olivia’s freebooting uncle, who has become locked in a ferocious power struggle with Malvolio, her ill-tempered steward. Yet it’s only with the arrival of an unsuspecting and very much alive Sebastian that events truly spiral out of control.

“Twelfth Night is very, very funny, but it’s also bittersweet,” says Schvey. “It's one of Shakespeare’s most satisfying comedies because it deals with fundamental and profound issues. It’s about falling in love, as all his comedies are, but also illusion, narcissism and delirium. It's not a play you laugh at and forget.”

Drummond, who serves as assistant director, points out that Malvolio’s “box tree” soliloquy, in which he unknowingly divulges his feelings for Olivia to a group of delighted rivals, has been called “the funniest scene in Shakespeare, and it can be played that way.

“But there’s also something uncomfortable about it,” Drummond says. “Malvolio reveals his deepest desires while the audience laughs at these characters who are making fun of him.

“It’s wildly funny, but also deeply sad.”

Daniel Hodges as Feste.

Schvey, who founded the PAD’s Globe summer program, notes that the intimate Hotchner Studio, which seats about 100, fosters an historically appropriate sense of give-and-take between cast and audience.

“One of the keys to working at the Globe is the concept of shared light,” Schvey says. “Plays were done during the day. Audience and actors were not divided the way they are in modern theatres.

“In the Hotchner, of course, we do use lighting effects, but there are ways to incorporate interaction between actor and audience that cannot be done in a larger space,” he adds. For instance, though Schvey's Twelfth Night transposes the story to 1950s Cuba, the sets remain minimal and the audience surrounds the stage on three sides.

“When an actor delivers a soliloquy, they speak directly to the audience, which is really closer to the heart of Shakespeare’s original practice,” Schvey says.

“The audience becomes a character in the play.”

Will Jacobs as Malvolio.

The cast of 19 is led by junior Kiki Milner as Viola, junior Anna Richards as Olivia and senior Will Jacobs as Malvolio. Sophomore Ricki Pettinato. and senior Louisa Kornblatt are Duke Orsino and Maria. Freshman Danny Washaleski is Sebastian.

Also featured are senior Eric Gustafson as Sir Toby Belch, sophomore Kate Needham as Sir Andrew Auguecheek and graduate student Daniel Hodges as Feste. Senior Ariel Saul is Fabian. Junior Mitchell Manar is Antonio.

Freshman Zack Schultz is the sea captain. Junior Clare Mulligan is the priest. Rounding out the ensemble are Katie Jeanneret, Jack Ritten, Cassie Roberts, Kilian Suchocki and Emma Quirk-Durben.

Scenic and costume design are by senior Quinlan Maggio and junior Maxine Wright, respectively. Henry Claude, the PAD’s music director for dance, based original compositions on 16th century melodies.

Choreography is by Christine Knoblauch-O’Neal, professor of the practice in dance. Sound and lighting are by junior David Levitt and Sean Savoie, design and technical coordinator. Props master is Emily Frei.

Assistant directors are Drummond and graduate student Jim Short. Sophomore Abby Mros is stage manager.

Twelfth Night begins at 8 p.m. Thursday and Friday, April 17 and 18; at 2 p.m. and 8 p.m. Saturday, April 19; and at 2 p.m. Sunday, April 20. Performances take place in the A.E. Hotchner Studio Theatre, located in the Mallinckrodt Center, 6465 Forsyth Blvd.

Tickets are $15, or $10 for students, seniors and Washington University faculty and staff, and are available through the Edison Theatre Box Office.

For more information, call (314) 935-6543.

Ron Freiwald

Shoulder to shoulder stand WUSTL undergraduate mathematicians: (from left) junior Alan Talmage; seniors Stephen Rong and Jason Zhang; junior Yu Tao Li; and sophomores Fangzhou Xiao and Anthony Grebe.

The university fielded 19 students in the competition, held in December. More than 4,000 students from 557 colleges and universities across the United States and Canada took part.

Three students must be designated in advance as the school team, and the team score is based on the three individuals' scores.

This year, the WUSTL team, consisting of sophomore Anthony Grebe, junior Patrick Lopatto and freshman Jongwhan Park, placed 10th out of  430 teams.

During the past 20 years, the WUSTL team has ranked among the top 10 teams seven times (and twice has made the top five).

The competition consists of two three-hour sessions during which contestants work individually on 12 problems.

"The Putnam is difficult," said Grebe, who was on both the Putnam and the Missouri Collegiate Mathematics Competition teams this year. "The median score is usually zero or some small fraction of a problem. Even if you solve only one problem completely, you're above average."

“Putman problems are written by absolute masters of problem writing," said junior Alan Talmage, who also sat for both competitions. "The problems usually have a very clever solution built into them. There will be some method that makes the problem easy, but finding that method will be very hard." 

Carl Bender, PhD, professor of physics and one of the Putnam coaches, gave the following example of an "easy" Putnam-style problem: Two mathematicians named A and B meet on the street. 'I haven't seen you for a while,' says A. 'How old are your three kids now?' 'Well,' says B, 'the product of their ages is 36.' 'Ach, tell me more,' says A. 'Oh, sorry,' says B. 'I should add that the sum of their ages is equal to the number of windows in the house over there.' 'I'm still at a loss.' says A. 'You've only told me two things and there are three kids.' 'My fault entirely," says B. 'I meant to tell you, the oldest is playing piano.' 'Oh, yes, of course, says A. Now I have it.'

"When you first read a Putnam problem, you don't have a clue what it is about," Bender said. "You don't know how many windows there are in that house, so how does that help? And what does playing piano have to do with it?"

"Normally, you spend the first hour of the competition poking around the problems, trying to understand them, but not doing very much in the way of computation," Talmage said. "Then, suddenly, you see something that looks interesting and you realize, 'Oh, I get it, if  I just do this, it will all fall out nicely.' "

Both Grebe and Talmage admit that the first phase of blind probing can be nerve-wracking. It's hard to tell whether your attempt isn't working or if you've just not gotten far enough, Grebe said.

"Should you give up on a problem and work on another one, or stick with same problem in the hope that in another hour you might get somewhere?"  he said.

"You get used to it," Talmage said. "But there are moments when you realize, OK, I have no idea what I'm doing. I'm just going to stop, clear my head, and look at the problem in a completely different way." 

What he says next may sound like a non sequitur but is actually typical of the contestants.

"I absolutely love it. I love doing math. I come out of the competition completely drained and normally just go straight home and fall asleep. But I absolutely love it." 

"Solving problems like these can be very habit forming," said Bender. At first you're in darkness, completely lost and without a clue," said Bender. "And then, suddenly, you see the point in a flash of inspiration. It is very rewarding."

Elizabeth Lowell Putnam established the William Lowell Putnam Intercollegiate Memorial Fund in memory of her husband, who graduated in mathematics from Harvard University. The first competition was held in 1938, and the Mathematical Association of America has sponsored the contest since then.

Students prepare for the Putnam during Friday afternoon practice sessions in the fall semester. The coaches this year were Xiang Tang, PhD, associate professor of mathematics; Richard Rochberg, PhD, professor of mathematics; and Bender. The practices featured free pizza, paid for by the Department of Mathematics from money won by past Putnam teams. (The top team can win as much as $25,000.)

An archive of Putnam problems can be found here.

Missouri Collegiate Mathematics Competition

WUSTL math students also did well in the 19th annual Missouri Collegiate Mathematics Competition, held March 27-28 at Saint Louis University. Forty-seven teams from colleges and universities across Missouri took part.

The two WUSTL teams took first and third place. A team consisting of Grebe, Talmage and sophomore Fangzhou Xiao captured first place with a perfect score of 100 — the first in the competition's history. The second team, consisting of junior Yu Tao Li and seniors Stephen Rong and Jason Zhang, took third place, with a score of 98.

The competition consists of two sessions, during each of which teams work collaboratively on five problems for 2 1/2 hours. The Missouri section of the Mathematical Association of America sponsors it. Since it began in 1996, a WUSTL team has captured first place 12 times.

Ron Freiwald, PhD, professor of mathematics and director of undergraduate studies in mathematics, and Blake Thornton, PhD, coordinator of lower-division teaching, shepherded the students through the state competition.

To view problems from this year's state competition, visit the competition’s website.

Fazzari

Steven Fazzari, PhD, a leading economics scholar known for an unwavering dedication to student mentoring in his three decades on the university faculty, will be installed Monday, April 21, as the inaugural Bert A. and Jeanette L. Lynch Distinguished Professor.

The Lynch estate, built on a successful box and packaging business, will also endow professorships named in honor of two of the most notable economics professors in university history, Douglass C. North, PhD, and the late Murray L. Weidenbaum, PhD.

North, the Spencer T. Olin Professor Emeritus in Arts & Sciences, joined WUSTL in 1983 as a Luce Professor. The co-recipient of the 1993 Nobel Memorial Prize in Economic Sciences is a pioneer in the field of institutional economics and a dedicated teacher. In addition to his teaching and research, North established the Center for New Institutional Social Sciences at WUSTL in 2001.

Weidenbaum, a longstanding authority on economic policy and government regulation who served as chairman of President Ronald Reagan’s first Council of Economic Advisers, died in March at age 87. Weidenbaum also was a veteran of the Department of Economics. In 1975, he founded the Center for the Study of American Business at WUSTL; in 2001, the center was renamed the Weidenbaum Center on the Economy, Government, and Public Policy; it became nationally recognized as an independent center that supports social-science research and organizes public outreach programs.

“It is deeply gratifying to honor the significant work and legacies of Doug North and Murray Weidenbaum through these professorships, as well as recognize Steve Fazzari’s important contributions to the profession,” Chancellor Mark S. Wrighton said.

“We are grateful to Bert and Jeanette Lynch for their very generous gift, which helps advance the study of economics at Washington University and provides our students access to some of the world’s very best scholars and teachers.”

Bert and Jeanette Lynch

A native of Blytheville, Ark., Bert Lynch moved to St. Louis to attend WUSTL, where he earned a bachelor’s degree in 1935 and a master’s degree in 1936, both in Arts & Sciences.

Native St. Louisan Jeanette Lichtenstein Lynch graduated from Mary Institute and attended WUSTL from 1933-34 before earning her degree in 1937 from Rollins College in Winter Park, Fla.

After their marriage, Bert Lynch joined his father-in-law’s firm, Superior Folding Box Co., eventually taking over the operation. He sold the business to a Kansas City-based box company in the early 1980s. He died in 1997; his wife died in 2013.

“This gift significantly raises the already strong reputation of Washington University’s Department of Economics and will allow the department to continue to attract world-class faculty,” said Barbara A. Schaal, PhD, dean of the Faculty of Arts & Sciences and the Mary-Dell Chilton Distinguished Professor. “Thanks to Bert and Jeanette Lynch, our future economics students will be exposed to some of the greatest minds in the field.”

The Lynch-generated professorships bring to 14 the number of endowed professorships in the Department of Economics.

The Douglass C. North Distinguished Professorship and the Murray L. Weidenbaum Distinguished Professorship will help the university attract and retain top faculty in the field of economics. Appointments to these new professorships will be made later.

Steven Fazzari

In addition to his teaching and research, in 2008 Fazzari took on a new leadership role as associate director of the Weidenbaum Center. There, he helps oversee the generation of scholarly research and the development of public affairs programs and other activities that address some of the most pressing public policy issues facing America.

His research explores two main areas: the link between macroeconomic activity and finance, particularly the financial determinants of investment spending; and the foundations of Keynesian macroeconomics.

Known as a gifted and popular teacher with several awards to his credit, Fazzari often collaborates with his students — past and present — on research projects and publications.

One such initiative is an innovative online resource called the “Muddy Water Macro,” a website where Fazzari and a team of students present well-organized, thoughtful, and broadly accessible ideas that they believe are central to understanding Keynesian economics. (The site can be accessed here).

Born in Racine, Wis., Fazzari spent his college summers working in a local foundry. His scholarly beliefs were informed by his middle-class Midwestern sensibility, and this draws him to collaborate with other faculty, such as Mark R. Rank, PhD, the Herbert S. Hadley Professor of Social Welfare. They co-teach a course, “Economic Realities of the American Dream,” which examines the American Dream’s historical meaning, the traditional pathways to achieving it, and its viability for the future.

Fazzari said his need to truly understand macroeconomic theories for teaching sparked his research interests.

“My connection with students has meant a lot to me, and many of my research ideas have emerged from my teaching,” he said.

He stays connected to his former students as well. One of his longest research collaborations has been with Barry Cynamon (AB '05), who is now a visiting scholar with the St. Louis Federal Reserve.

Most recently, they co-edited “After the Great Recession: The Struggle for Economic Recovery and Growth,” published in 2013, as well as a recent research paper widely cited in the national news providing solid evidence for why the U.S. economy has remained slow growing.

Cynamon explained the genesis of their collaboration:

“A conversation that began during office hours while I was taking Steve’s undergraduate macro seminar continued through graduate school, several research papers, an edited volume, and enough miles of walking and talking to keep us both in reasonably good shape.”

Fazzari continues to teach a wide range of macroeconomic courses, from introductory freshman classes to advanced doctoral seminars. His teaching has been recognized with the Governor’s Award for Excellence in Teaching, the Excellence in Teaching Award from Emerson Electric Co., and WUSTL's Distinguished Faculty Award in 2007.

Tinianow

Alex Tinianow has been selected to receive the 2014 Harrison D. Stalker Award from the Department of Biology in Arts & Sciences at Washington University in St. Louis. 

This award is named in honor of the late Harrison D. Stalker, PhD, who was a member of the biology faculty from 1942 to 1982. In addition to being a renowned evolutionary biologist and an inspiring teacher, Stalker was a great advocate of the arts and a world-class photographer. 

The award is given annually to a graduating biology major whose undergraduate career combines outstanding scientific scholarship with significant contributions in the arts and humanities.

Tinianow, a biology major with a neuroscience track and a jazz studies minor, is a summa cum laude candidate who exemplifies the combination of scientific contribution and artistic expression this award seeks to recognize.

Tinianow translated his interest in how the brain works into a two-year undergraduate research project with Erik Musiek, MD, PhD, in the David Holtzman laboratory in the Department of Neurology at the School of Medicine. There, he examined how chronic jetlag influences the progression of Alzheimer’s disease in mouse models. Jetlagged mice, whose light-dark schedule was advanced by six hours once a week for eight weeks, had more amyloid-beta plaque burden than non-jetlagged control mice.

In addition to working in the lab, Tinianow currently serves as a tutor for the introductory biology course, Principles of Biology I.

Tinianow combines scientific curiosity with achievements in the arts. A talented musician, he has played piano and alto saxophone in the jazz combo and jazz band programs in the Department of Music. He also helped to create a new vocal jazz ensemble that is now a permanent part of the jazz combo program, and he and the group have performed in the Jazz at Holmes Concert Series.

In addition, Tinianow is the artistic director and former assistant musical director and treasurer of The Ghost Lights, a student-run WUSTL a cappella group.

His interests in neuroscience and music also converged when he assisted in a clinical study at the School of Medicine, led by Gammon Earhart, PhD, that examined the use of tango to treat Parkinson’s disease patients, who often move better to music than they can in silence.

After graduation, Tinianow plans to work as a research assistant in preparation for applying to either a PhD or MD/PhD program in neuroscience.

Image courtesy of J. Ponsonby

Shorthorn bulls named St. John and Gaudy in this painting by Thomas Freebairn Wilson are products of human control over breeding. The fascination with livestock improvement in 19th-century Britain is reflected in portraits of prize animals like this one — enough of them to support a trade in itinerant livestock painting.

Many of our ideas about domestication derive from Charles Darwin, whose ideas in turn were strongly influenced by British animal-breeding practices during the 19th century, a period when landowners vigorously pursued systematic livestock improvement. 

It is from Darwin that we inherit the ideas that domestication involved isolation of captive animals from wild species and total human control over breeding and animal care.

But animal management in this industrial setting has been applied too broadly in time and space, said Fiona Marshall, PhD, professor of anthropology at Washington University in St. Louis. It is not representative of the practices of the Neolithic herders who first domesticated animals nor — for that matter — of contemporary herders in nonindustrial societies.

Together with Keith Dobney, PhD, of the University of Aberdeen in Scotland; Tim Denham, PhD, of the Australian National University; and José Capriles, PhD, of the Universidad de Tarapacá in Chile, Marshall wrote a review article that summarizes recent research on the domestication of large herbivores for "The Modern View of Domestication," a special feature of The Proceedings of the National Academy of Sciences published April 29.

Recent research on the domestication of donkeys, camelids (which includes dromedaries, Bactrian camels, llamas and alpacas) pigs, cattle, sheep and goats suggests that neither intentional breeding nor genetic isolation were as significant as traditionally thought, the scientists said.

“Our findings show little control of breeding, particularly of domestic females, and indicate long-term gene flow, or interbreeding, between managed and wild animal populations,” Marshall said.

Why is it important to get domestication right? “Our livestock is losing genetic diversity even faster than some wild animals, because of management practices like artificial insemination,” Marshall said. “We took only a bit of the diversity from the wild for domestication, and what we’re looking at now is lopping it off really fast so we’ll be left with little diversity to survive all the climate and disease issues we’re facing. It really is a crisis situation.

“If we don’t understand what it is we might be about to lose, then we don’t count the cost of loss accurately or know how to plan for the future,” she said.

A walk on the wild side

Stine Rossel/PNAS

One of the markers of domestication is a reduction in size, but archeological evidence indicates size decreases were slow and inconsistent. Donkeys buried 5,000 years ago in an early pharaonic mortuary complex (above) have proportions similar to those of the African wild ass, but the bones of domesticated donkeys found at another, much older site are significantly smaller than those of wild asses.

For most of history, artificial selection on large herbivores was probably weak, Marshall said. “Herders could not afford to kill many animals, particularly large-bodied animals with long gestation periods. To keep herd size stable, herders probably culled or castrated males surplus to the growth needs of the herd, allowing all females to breed,” she said. These management practices placed only light selection pressure on the herd’s gene pool.

Paradoxically, environmental selection may, in many instances, have been stronger than artificial selection. Early herds were vulnerable to disease, droughts and storms, disasters that would have forced pastoralists to replenish herds from wild populations better adapted to harsh local conditions.

Sometimes domesticated animals were intentionally bred with wild ones, Marshall said. “Wild animals are generally faster, stronger and better adapted to the local conditions than domesticated ones. So, for example, Beja herders in Northeastern Africa intentionally bred their donkeys with African wild asses in order to produce stronger transport animals."

“And sometimes interbreeding was accidental,” she said. “Even today in the Gobi, researchers report that domestic camels sometimes join wild herds after becoming separated from their own. Wild and domestic camels meet at shared oases, and wild males also can become extremely aggressive and may collect domestic females to the dismay of pastoralists.” 

J. Capriles

So confused is the genetic history of llamas that some are in fact chimeras; they have cells in their bodies from two distinct maternal lineages.

In the Andes, Capriles said, wild and domestic camelids have interbred in such complex ways that alpacas are maternally related to both wild vicunas and guanacos, and the same is true for llamas.

Artificial selection was probably weakest and gene flow highest in the case of pack animals such as donkeys or camelids. But even in the case of pigs or cattle, interbreeding between domestic and wild animals has created long and complex evolutionary and domestication histories that challenge assumptions regarding genetic isolation and long-held definitions of domestication.

The curl in the pigs’ tails

The domestication of pigs is one of these stories. Dobney, Greger Larson, PhD, and their team have shown that pigs were domesticated at least twice, in eastern Anatolia and in central China. Analysis of mitochondrial DNA (DNA in a cell organelle that is inherited from the mother) shows that early herders took pigs with them from Anatolia to western Europe. And analysis of ancient DNA shows that, once in Europe, the domesticated pigs interbred with the wild boars. These hybridized populations then rapidly replaced the original domesticates, first in Europe and then, later, across Anatolia itself.

In China, the story is somewhat different. There is little evidence that the domestic herds in central China interbred with wild boars. But early agriculturists took their pigs to southeastern Asia and there, deliberately or accidentally, recruited local wild boar lineages into their domestic stock.

All of the New Guinea domestic pigs and those of the islands in the tropical Pacific Ocean carry DNA from those southeast Asian wild boar populations.

The interesting question is why the pigs in central China didn’t interbreed with wild boar populations in central China. Dobney suggests that management practices may have made a difference. It is possible that in China where settlements were dense, people started keeping pigs in pens, whereas in Europe, even in medieval times, people took their pigs to forage in the forests, where they might encounter wild boars.

The pig story illustrates how much our understanding of domestication events has changed. The anomaly is the isolated domestic population, not the prolonged interbreeding among domestic and wild animals, which in most domesticated species seems to have continued to recent times.

What would Darwin say?

“The research is really exciting because it is making us completely rethink what it means to be domesticated,” Marshall said. “The boundaries between wild and domesticated animals were much more blurred for much longer than we had realized."

“To untangle the history of domestication,” Denham said, “scientists will need to bring to bear all of the evidence at their disposal, including archeological and ethnographic evidence, and the analysis of both modern and ancient DNA.”

“We must also investigate sources of selection more critically," Marshall said, "bearing in mind the complex interplay of human and environmental selection and the likelihood of long-term gene flow from the wild."

It’s probably fortunate the Darwin had clear examples of animal breeding to consider as he thought about evolution. The first chapter of "On the Origin of Species" discusses the domestication of animals such as as pigeons, cattle and dogs, and Darwin then uses artificial selection as a springboard to introduce the theory of natural selection.

It turns out that animal domestication is more complex, and the role of natural selection more important than Darwin thought. It is also the case that the people who first domesticated animals valued wild ones more than did Darwin’s Victorian neighbors. 

“The Modern View of Domestication,” a special issue of PNAS edited by Greger Larson and Dolores R. Piperno, resulted from a meeting entitled “Domestication as an Evolutionary Phenomenon: Expanding the Synthesis,” held April 7–11, 2011, that was funded and hosted by the National Evolutionary Synthesis Centre (National Science Foundation EF-0905606) in 2011.

Donald Knuth/Ethnobotanical gardens/CC License

The changes that took place as this plant was domesticated were so dramatic that its ancestry became a mystery for many decades. The plant is teosinte, now known to be the progenitor of maize (or corn, as it is called in the United States).

“The Modern View of Domestication,” a special feature of The Proceedings of the National Academy of Sciences (PNAS) published April 29, raises a number of startling questions about a transition in our deep history that most of us take for granted. At the end of the last Ice Age, people in many spots around the globe shifted from hunting animals and gathering fruits and tubers to cultivating livestock and plants.

It seems so straightforward and yet the more scientists learn, the more complex the story becomes. Recently, geneticists and archeologists working on domestication compared notes and up popped a question of timing. Did domesticating a plant typically take a few hundred or many thousands of years?

Genetic studies often indicate that domestication traits have a fairly simple genetic basis, which should facilitate their rapid evolution under selection. On the other hand, recent archeological studies of crop domestication have suggested a relatively slow spread and fixation of domestication traits.

In this special issue of PNAS, Washington University in St. Louis biologist Ken Olsen, PhD, and colleagues ask whether complex genetic interactions might have slowed the rate at which early farmers were able to shape plant characteristics, thus reconciling the genetic and archeological findings.

Olsen, associate professor in the Department of Biology in Arts & Sciences, together with colleagues from Oklahoma State University and the University of Guelph in Ontario, Canada, conclude that these interactions are not a key factor in domesticated plants. The process of domestication, Olsen said, favored gene variants (alleles) that are relatively insensitive to background effects and highly responsive to selection.

But finding these alleles in the first place must have difficult, Olsen said. Only a subset of the genes in the wild population would have reliably produced a favored trait regardless of the crop variety into which they were bred and regardless of where that crop was grown. So the early stages of domestication might have been beset by setbacks and incomprehensible failures that might help explain the lag in the archeological record.

“What we are learning suggests there’s a whole lot of diversity out there in wild relatives of crop plants or even in landraces, varieties of plants and animals that are highly adapted to local conditions," Olsen said, "that wasn’t tapped during the domestication process.”

“These plant populations could provide the diversity for continued breeding that is going to be very important as the world faces climatic change," he said. "This is why it is important we understand the early stages of domestication.”

Two possible speed bumps

Many crops are distinguished from their wild ancestors with a suite of traits called the domestication syndrome. This includes seeds that remain attached to the plant for harvesting (a trait called nonshattering), reduced branching and robust growth of the central stem and bigger fruits, seeds or tubers.

Stronglk7/cc License

Like other domesticated cereals, foxtail millet has nonshattering spikes that retain their seeds during harvesting.

Over the past 20 years, researchers have begun to identify the genes that control some of the most important domestication traits, no easy task in the days before rapid sequencing, because they had to start with plant traits and work back to unknown genes.

Erikeltic/English wikipedia/cC License

Coat color in Labrador retrievers is due primarily to epistatic interactions between two genes. One gene determines whether the dog has a dark or yellow coat. The second gene determines whether the darker dogs will be black or chocolate labs (and whether yellow labs have brown or pink noses).

This work showed that many domestication traits were under the control of single genes. For example the gene teosinte branched1 (tb1) converts highly branched teosinte plants into single stalks of corn.

But the seeming importance of single genes could have been an artifact of the method used to identify domestication genes, which required the researcher to pick “candidate” genes and, perhaps, prematurely narrow the search, overlooking indirect genetic effects.

“Little is known about the underlying genetics of domestication,” Olsen said. “We decided to look at genetic mechanisms for modifying plant phenotypes that hadn’t been explored before, in part because not much data is available.”

The new work examines the possibility that two indirect effects — the influence of the genetic background on the expression of a gene (called epistasis) and the effects of the environment on the expression of genes — might have slowed the selection of plants with the desired traits.

Epistasis and environmental effects in domestication genes

By selecting animals for coat color, animal breeders may have stabilized certain epistatic and environmental interactions in companion animals (see photos at right). But when the plant scientists looked at comparable genetic mechanisms in domesticated plants, they found the reverse to be true. Farmers seem to have selected for plant variants that were insensitive to epistatic and environmental interactions.

Thaifong/CC License

A good example of the influence of the environment on genes is the coat pattern in a seal-point Siamese cat. Siamese cats have a heat-sensitive enzyme that is inactivated by body heat on the head and torso of the cat, leaving the coat there uncolored, but is active on the cooler ears, face and tail, turning these areas dark.

Shattering in domesticated foxtail millet provides an example of insensitivity to epistasis. Branching in maize illustrates insensitivity to environmental effects.
Shattering in foxtail millet and its wild ancestor, green millet, is controlled by two stretches of DNA containing or linked to genes that underlie this trait, a major one called QTL 1 and a minor one called QTL2. In this as in other epistatic interactions, the effect of an allele at one location depends on the state of the allele at the other location. But when wild and domesticated plants are crossed, these “genetic background effects” are not symmetric.

Shattering in plants with a wild green-millet allele at the QTLI location depends on the allele at the QTL2 location. In contrast, shattering in plants with the foxtail-millet allele at QTL1 is unaffected by the allele at the QTL2 location.

In the limited number of examples at their disposal, the scientists found it to be generally true that that domesticated alleles were less sensitive to genetic background than wild alleles. The domestication genes, in other words, tended to be ones that would produce the same result even if they were introduced into a different crop variety.

Teosinte provides a good example of the sensitivity of gene expression to the environment. Teosinte is strongly affected by crowding. When a teosinte plant with a wild tb1 gene is repeatedly backcrossed with maize, it produces highly branched plants in uncrowded growing conditions but plants with smaller lateral branches when it is crowded.

Again, however, the effect is not symmetric. The domesticated trait is less sensitive to the environment than the wild trait; plants with the domesticated tb1 gene allele are unbranched whether or not they are crowded.

Unlike companion-animal breeders, early farmers seem to have selected domestication-gene alleles that are insensitive to genetic background and to the environment. This process would have been slow, unrewarding and difficult to understand, because the effects of gene variants on the plant weren’t stable. But once sensitive alleles had been replaced with robust ones, breeders would have been able to exert strong selection pressure on plant traits, shaping them much more easily than before, and the pace of domestication would have picked up.

No wonder the archeological record indicates there were false starts, failed efforts and long delays.

"The Modern View of Domestication," a special issue of PNAS edited by Greger Larson and Dolores R. Piperno, resulted from a meeting entitled “Domestication as an Evolutionary Phenomenon: Expanding the Synthesis,” held April 7–11, 2011, that was funded and hosted by the National Evolutionary Synthesis Centre (National Science Foundation EF-0905606) in 2011.

Jerry Nauheim Jr./WUSTL Photos

Petra Levin (left) and Joseph Jez, pictured in the Department of Biology's greenhouse on the Danforth Campus, are co-directors of the graduate program in Plant and Microbial Biosciences at Washington University in St. Louis. A generous gift from William H. Danforth is expanding the number of fellowships in the program.

Washington University in St. Louis Chancellor Mark S. Wrighton has announced the creation of new four-year Danforth Fellowships for students in any PhD program with an interest in plant sciences. The new fellowships will enhance the St. Louis regional effort in plant sciences and will contribute to building the collaboration between Washington University and the Donald Danforth Plant Science Center.

At the March 7 Board of Trustees meeting, the following Washington University in St. Louis faculty members were appointed with tenure, promoted with tenure or granted tenure, effective July 1 unless otherwise noted.

Appointment with tenure

Shabaana Abdul Khader, PhD, as associate professor of molecular microbiology (effective March 7); 

Todd P. Margolis, MD, PhD, as professor of ophthalmology and visual sciences (effective March 7); and

Anya Plutynski, PhD, as associate professor of philosophy (effective March 7).

Promotion with tenure

Catherine S. Adcock, PhD, to associate professor of history;

J. Dillon Brown, PhD, to associate professor of English;

William S. Bubelis, PhD, to associate professor of classics;

Bruce A. Carlson, PhD, to associate professor of biology;

David A. Fike, PhD, to associate professor of earth and planetary sciences;

Elizabeth S. Haswell, PhD, to associate professor of biology;

Crickette Sanz, PhD, to associate professor of anthropology;

Monika J. Weiss to associate professor of art;

Kilian Q. Weinberger, PhD, to associate professor of computer science and engineering;

Timothy M. Miller, MD, PhD, to associate professor of neurology (effective March 7);

Alexander S. Krupnick, MD, to associate professor of cardiothoracic surgery (effective March 7);

David D. Limbrick Jr., MD, PhD, to associate professor of neurological surgery (effective March 7);

Daniel S. Marcus, PhD, to associate professor of radiology (effective March 7);

Erika L. Pearce, PhD, to associate professor of pathology and immunology (effective March 7);

Zhude Tu, PhD, to associate professor of radiology (effective March 7);

Granting of tenure

Markus Baer, PhD, as associate professor of organizational behavior;

Xiumin X. Martin, PhD, as associate professor of accounting (effective April 1); and

Lamar Pierce, PhD, as associate professor of strategy.

Sid Hastings/WUSTL Photo Services

Washington University biologist Ken Olsen, who studies the genetic basis of evolution in plants, and archeologist Fiona Marshall, whose research focuses on animal domestication in Africa, enjoy an interdisciplinary chat.

We all think we have a rough idea of what happened 12,000 years ago when people at several different spots around the globe brought plants under cultivation and domesticated animals for transport, food or fiber. But how much do we really know?

Recent research suggests less than we think. For example, why did people domesticate a mere dozen or so of the roughly 200,000 species of wild flowering plants? And why only about five of the 148 species of large wild mammalian herbivores or omnivores? And while we’re at it, why haven’t more species of either plants or animals been domesticated in modern times?

If nothing else, the tiny percentages of domesticates suggests there are limitations to human agency, and that it almost certainly is not true that people can step in and completely remodel through artificial selection an organism shaped for millennia by natural selection. 

The small number of domesticates is just one of many questions raised in a special issue of the Proceedings of the National Academy of Sciences published online April 21. 

The issue is the product of a 2011 meeting of scholars with an interest in domestication at the National Evolutionary Synthesis Center, a nonprofit science center jointly operated by Duke University, the University of North Carolina at Chapel Hill and North Carolina State University.

Of the 25 scholars at the conference, two were from Washington University in St. Louis: Arts & Sciences’ Fiona Marshall, PhD, professor of archaeology, who studies animal domestication, and Kenneth Olsen, PhD, associate professor of biology, who studies plant domestication.

Both Marshall and Olsen are currently engaged in research on the crumbling margins of domestication where questions about this evolutionary process loom the largest. 

Marshall studies two species that are famously ambivalently domesticated: donkeys and cats. Olsen studies rice and cassava and is currently interested in rice mimics, weeds that look enough like rice that they fly under the radar even when rice fields are handweeded.

Both Marshall and Olsen contributed articles to the special PNAS issue (see The story of animal domestication retold and Genetic study tackles mystery of slow plant domestications) and helped write the introductory essay that raises the big questions confronting the field.

“This workshop was especially fun,” said Olsen, “because it brought together people working on plants and animals and archeologists and geneticists. I hadn’t really thought much about animal domestication because I work primarily with plants, so it was exciting to see the same problem from a very different perspective.” 

How much of it was our doing?
Many of our ideas about domestication are derived from modern experience with animal breeding. Anyone familiar with the huge variety of dog breeds, all of which belong to the same subspecies of the gray wolf, has some appreciation of the power of selective breeding to alter appearance and behavior.

Brian Hare/Duke University

Perhaps the most famous experiment in domestication is a project in Russia that turned silver morphs of the wild red fox into tamer and more dog-like silver foxes in just 40 generations. But the silver foxes were kept in cages on a fox farm where they were sheltered and fed and illicit liaisons with wild foxes were thwarted. How representative was this experiment of prehistoric domestication events?

But what about self-fertilizing or wind-pollinated plants, or for that matter, domesticated animals accidentally or deliberately bred with wild relatives?

Recent evidence that cereal crops, such as wheat or barley, evolved domestication traits much more slowly than had been thought has led to renewed interest in the idea that selection during domestication may have been partly accidental.

Charles Darwin himself drew a distinction between conscious selection, in which humans directly select for desirable traits, and unconscious selection, where traits evolve as a byproduct of natural selection in crop fields or from selection on other traits.

“The big focus right now is how much unintentional change people were causing environmentally that resulted in natural selection altering both plants and animals,” said Marshall.

“We used to think cats and dogs were real outliers in the animal domestication process because they were attracted to human settlements for food and in some sense domesticated themselves. But new research is showing that other domesticated animals may be more like cats and dogs than we thought. 

Out of copyright/Creative Commons license

Why weren’t zebras ever domesticated? Baron Rothschild frequently drove a carriage pulled by zebras through the streets of 19th-century London. In “Guns Germs and Steel,” Jared Diamond says the reason zebras were not domesticated is that they are extraordinarily vicious and will bite and not let go. But why weren’t people able to modify this temperament if they were able to gentle wolves into dogs?

“Once animals such as donkeys or cattle were caught,” Marshall said, “the changes humans sought to make were pretty minimal. Really it just came down to culling a few of the males and breeding all of the females.” 

Even today, she points out, African pastoralists can afford to kill only four out of every 100 cows or they run the risk that drought and disease will wipe out the entire herd. “So I think outside of industrialized societies or special situations, artificial selection was very weak,” she said.

“In the donkeys and other transport animals, it’s not affiliative [tame] behavior the herders want,” Marshall said. “What they care about more than anything else is that their animals stay alive.” 

So artificial selection is acting in the same direction as natural selection, or maybe pushing even harder, because humans often place animals in harsher conditions than natural ones.

“The comparable idea for plants,” said Olsen, “is the dump heap hypothesis, originally proposed by Edgar Anderson, a botany professor here at Washington University. The idea is that when people threw out the refuse of plant foods, including seeds, some grew and again set seed, and in this way people inadvertently selected species they were eating that also did well in the disturbed and nutrient-rich environment of the dump heap.”

“Cultivation practices play a huge role in selection,” said Olsen. “Traditionally in Southeast Asia, many different varieties of rice were grown simultaneously in a given field. It was a bet-hedging strategy,” he said, “that ensured some plants would survive and produce seed even in a bad season.” So it wasn’t people selecting the crop plants directly so much as people changing the landscape in ways that altered the selection pressure on plants. 

How best to time travel
Questions about the original domestication events are difficult to answer because plants and animals were domesticated before humans invented writing, and so figuring out what happened has been a matter of making do with the limited evidence that has survived.

The problem is particularly difficult for animal domestication because what matters most is animal behavior, which leaves few traces. In the past, scientists tried measuring bones or examining teeth, looking for age or size differences or pathology that might plausibly be related to animals living with people. 

“Sometimes there aren’t morphological shifts that are easy to find or they’re too late to tell us anything,” Marshall said. “We’ve gone away from morphological identifiers of domestication, and we’re going with behavior now, however we can get it. If we’ve got concentrations of dung, that means animals were being corralled,” she said.

Olsen, on the other hand, seeks to identify genes in modern crop species that are associated with domestication traits in the plant, such as an erect rather than a sprawling architecture. The techniques used to isolate these genes are difficult and time consuming and may not always penetrate as deeply into the past as scientists had once assumed because present-day plants are only a subset of the crop varieties that may have once existed.

Prof. SAXX/Wikimedia Commons

Aurochs, the ancestors of modern cattle, depicted in this cave painting in Lascaux, France, are now extinct. The last recorded auroch died in Poland in 1627. Marshall worries that the erosion of genetic diversity symbolized by this extinction might make it harder to remold domesticated species to meet the challenges of climate change.

So both Marshall and Olsen are excited by recent successes in sequencing ancient DNA. Ancient DNA, they say, will allow hypotheses about domestication to be tested over the entire evolutionary time period of domestication.

Another only recently appreciated clue to plant domestication is the presence of enriched soils, created through human activities. One example is the terra preta in the Amazon basin, which bears silent witness to the presence of a pre-Columbian agricultural society in what had been thought to be untouched forest. 

By mapping distributions of enriched soils, scientists hope to better understand how ancient people altered landscapes and the effects that had on plant communities.

“It is really clear,” Marshall said, “that we need all the different approaches that we can possibly get in order to triangulate back. We’re using all kinds of ways, coarse-grained and fine, long-term and short, because the practical implications for us are quite great.”

After all, the first domestications may have been triggered by climate change at the end of the last ice age — in combination with social issues. 

As a result, people abandoned the hunter-gatherer lifestyle they had successfully followed for 95 percent of human history and turned instead to the new strategies of farming and herding. 

As we head into a new era of climate change, Marshall said it would be comforting to know that we understood what happened then and why.

“The Modern View of Domestication,” a special issue of PNAS edited by Greger Larson and Dolores R. Piperno, resulted from a meeting titled “Domestication as an Evolutionary Phenomenon: Expanding the Synthesis,” held April 7–11, 2011, that was funded and hosted by the National Evolutionary Synthesis Centre (National Science Foundation EF-0905606) in 2011.

Courtesy Photo

Smith

“It is an intense day,” said Smith, a student of anthropology, public health and women, gender and sexuality studies. “But I want to practice what I am learning. I view all of these experiences as completing, extending and challenging the education. I’ve learned so, so much.”

Smith is one of seven recipients of the 2014 Gerry and Bob Virgil Ethic of Service Award. The annual honor is given to members of the WUSTL community who have donated their time and resources to improving the St. Louis region.

To learn more about the other award winners honored this week, visit here.

Jami Ake, assistant dean in the College of Arts & Sciences, said Smith’s work as court advocate with the St. Louis County Domestic Violence Court and as a facilitator at RAVEN, a batterer intervention program, demonstrates Smith’s willingness to tackle society's toughest problems.

WHITnEy Curtis/WUSTL Photos

Winners of ths year's Gerry and Bob Virgil Ethic of Service Award gather. In back, from left, are: freshman Daniel Giuffra, alumna Aleeza Granote (MSW '04), Mike Pereira (University College, Class of 2015), Maxine Clark (Board of Trustees member), and senior Jennifer Rowley, In front, from left, are: senior Alaina Smith, Bob Virgil, Gerry Virgil and Patrick Gibbons (professor emeritus of physics).

“Alaina is one of those tremendously intellectually engaged students who is not at all satisfied with the theoretical answers to problems or with learning that remains solely within the boundaries of a classroom,” Ake wrote in her nomination. “It has been her work outside the classroom — or more precisely, her way of bringing the classroom into the community and the community into the classroom — that has made such a profound impact on both the Washington University community and St. Louis.”

At the county court, Smith helps petitioners file orders of protection against partners or stalkers and connects them to community resources. Most importantly, Smith listens.

“I always start by asking, ‘How are you doing?’” Smith said. “Sometimes they just need to talk about how they are afraid and unsure. And often they are angry at how surprisingly complicated the process can be.”

Smith knows it’s easy to make assumptions about her clients — that they are weak or somehow to blame for their troubles. But Smith has learned these situations are complicated. Take, for instance, the woman who chose to drop her order of protection against her ex-husband. She wanted to be better prepared in case he retaliated violently.  So Smith worked with her to devise an emergency safety plan.

“She told me she always sleeps on the couch so she could protect her kids if he should come in,” Smith said. “So we discussed what room she would go to if that happened. You want to avoid the kitchen because there are knives. You want to avoid the bathroom because there are hard surfaces. You want to think about all of these things, like: Do your children know how to call 911? I would never assume I know what’s best for them. These women are incredibly smart and strategic, often for the sake of their children.”

At RAVEN, Smith helps the men who cause such fear. She co-facilitates a weekly group for seven men, most of whom the court ordered to attend. Through education and discussion, RAVEN tries to undo a lifetime of destructive habits and beliefs.

“We teach that we always have control over our actions — that anger is an emotion, but violence is a choice,” Smith said. “It sounds strange, but we empower the men to make healthier decisions. Needless to say, not everyone wants to be there, and it takes time for these messages to sink in.”

The program typically takes 17 months to complete. Many drop out. But others change.

“When many of the men start, they will say things like, ‘She started it,’ or ‘I didn’t hit her that hard,’ or ‘The kids weren’t around,’” Smith said. “At the end, they aren’t making excuses. They know it doesn’t matter what she was doing; they own what they’ve done.”

The work is challenging but not dangerous, Smith said. A co-facilitator is always present, and the men typically are on their best behavior, she said.

“People are scared for me a lot, but the bigger battle is to get the men to drop their facade of, ‘I’m cool, calm and collected,’ and get them to be honest,” Smith said. “There have been times when I have felt disrespected, but those cases are more the exception. I believe truly that people, at their core, are good.”

Originally from the Boston area, Smith has adopted St. Louis as her home. Since her arrival as a freshman, she has volunteered in the community. She has provided child care at the Women’s Safe House, tutored elementary school students at Niños Youth Mentoring and fostered cats for Stray Rescue.

On campus, she has worked for the Community Service Office, staged“The Vagina Monologues” and served as co-president of GlobeMed, spearheading the group’s partnership with a grassroots nongovernmental organization in Uganda.

Smith plans to stay in St. Louis after graduation and would like to work in a communications or outreach capacity at a nonprofit service agency.

“I love this city and have some deep roots here now,” Smith said. “When I came to Wash. U., I was interested in learning about the structural inequalities in other countries. But now I see those things exist in our own communities. I just feel so grateful for all of the mentors who have given me the opportunities to get involved right here.”

Technology is revolutionizing more than just how we shop, communicate and entertain ourselves. It is also changing how humanists analyze texts in a growing field called the digital humanities.

Now, scholars of literature and history can take thousands of digitized texts and use a variety of computational tools to engage in what some have called distant reading, a supplement to the close analyses that long have formed the basis of literary criticism or historical inquiry.

They can track word usage, spelling changes, publication patterns, the rise of a genre or the development of an idea. Programs even analyze an author’s style based on, for example, the number of times he or she uses a certain word.

SID Hastings/WUSTL Photos

Joseph Loewenstein, PhD, director of the Humanities Digital Workshop and the Interdisciplinary Project in the Humanities, is leading a “vertical seminar” to help senior and junior faculty, postdoctoral students, graduate students and staff better understand the digital humanities and use its techniques.

Scholars of all types are starting to use these tools. Joseph Loewenstein, PhD, director of the Humanities Digital Workshop and the Interdisciplinary Project in the Humanities at Washington University in St. Louis, is leading a “vertical seminar” to help senior and junior faculty, postdoctoral students, graduate students and staff better understand the digital humanities and use its techniques.

“[Digital analysis] changes the way a humanist works,” said Loewenstein, a professor in the Department of English in Arts & Sciences. “Most of the time, we work by ourselves, but nobody has enough expertise to do alone the kinds of projects that our tools make it possible for them to imagine. The innovative digital scholar has to collaborate.”

Participants in the Vertical Seminar in the Humanities come from a spectrum of disciplines, including German, comparative literature, American culture studies, African and African-American studies, history and English. Two staff members from the Humanities Digital Workshop complete the group.

“One of the goals of the vertical seminar was to get the expertise and obsessions of people at different career stages and from different disciplines talking to each other,” Loewenstein said.

The seminar is part of a three-year, $500,000 grant from The Andrew W. Mellon Foundation awarded to WUSTL in 2011 in support of five vertical seminars in the humanities. Steven Zwicker, PhD, the Stanley Elkin Professor in the Humanities and professor of English, applied for the grant and asked Loewenstein to teach this seminar.

Everyone in the course has a project. Lynne Tatlock, PhD, the Hortense and Tobias Lewin Distinguished Professor in the Humanities and director of the Comparative Literature program, is expanding on her book, “German Writing, American Reading: Women and the Import of Fiction 1866–1917.” The book explored what happened to translations of German texts when they reached America. How were they marketed? Who read them? How popular were they?

SID Hastings/WUSTL Photos

Lynne Tatlock, PhD, the Hortense and Tobias Lewin Distinguished Professor in the Humanities and director of the Comparative Literature program, leads a discussion during the seminar.

Tatlock used digital tools to keep track of all of the translations — she wrote about more than 100 books. But since she published the book, a database of circulation records from the late 1800s from a small public library in Muncie, Ind., has gone online. The library carried many translations of German books, and Tatlock is exploring how often they were checked out and by whom.

“I found out that even though my book was already in print, I still had a lot to do,” Tatlock said. “So that pulled me back into the digital humanities.

A collaboration with Matt Erlin, PhD, chair of the Department of Germanic Languages and Literatures, also fostered Tatlock’s work with digital humanities tools. The pair co-edited “Distant Readings: Topologies of German Culture in the Long Nineteenth Century,” a collection of essays that grapple with digesting the huge number of publications released in Germany in the 19th century.

Anupam Basu, PhD, WUSTL’s Mark Steinberg Weil Early Career Fellow in Digital Humanities, is also a member of the seminar. Basu, who always has been interested in computer science, but chose instead to study literature, learned about the digital humanities while working on his dissertation.

Sid Hastings/WUSTL Photos

Anupam Basu, PhD, the Mark Steinberg Weil Early Career Fellow in Digital Humanities, discusses his vertical seminar project as Joseph Loewenstein, PhD, listens.

“It was a really good way to complement and expand the horizons of the kind of research that I was already doing and other humanities scholars were doing,” Basu said.

He is particularly interested in Shakespeare and other Renaissance authors. Most of the books from this era have been digitized in Early English Books Online (EEBO-TCP), a collection of most of the printed works of England, Ireland, Wales, Scotland, and British North America from 1473 to 1700.

Because he already has advanced computer skills, Basu’s project for the seminar has been to develop tools to help scholars explore the EEBO-TCP database. Many of the tools that he and Steve Pentecost, senior digital humanities specialist, have created are online at earlyprint.wustl.edu.

For Basu, the seminar offers a chance for scholars to see their field or research question in a new light, by interacting with people from different disciplines and using new techniques to analyze familiar material.

“That wonderful dissociative alignment doesn’t happen in a normal seminar format where you don’t have such widely different perspectives,” Basu said.

“Sometimes you have someone discovering a new thing they can do that they haven’t thought of before, or that they didn’t know was possible before,” Basu said. “And in any sort of seminar or class, that’s the best thing you can have.”

Grant Wahl, senior writer at Sports Illustrated and an analyst for Fox Soccer, and Clark Hunt, chairman and CEO of the Kansas City Chiefs and a founding investor in Major League Soccer, are among the panelists. The discussion will be at 7 p.m. Monday, May 5, in 
Graham Chapel on the university’s Danforth Campus.

Dan Flynn, secretary general of the U.S. Soccer Federation, and Taylor Twellman, ESPN soccer analyst and the 2005 Major League Soccer Most Valuable Player, are also on the panel.

The panel discussion is free and open to the public.

Michael J. MacCambridge, adjunct instructor in communications and journalism in University College who teaches "Soccer & The Global Village: The Culture of the World Cup," will serve as moderator.

"Soccer has never been more popular in the United States than it is right now,” MacCambridge said. “This panel is an attempt to discuss the implications of that, not only for the game itself, but for the rest of the American sporting landscape.

"I'm thrilled to have such respected and fascinating guests joining the discussion. Clark, as both an NFL and MLS owner, is an ideal person to speak about soccer's growth within the larger context of American sport; Dan has been at U.S. Soccer for more than 15 years and has seen monumental changes in that time; Taylor brings a player's understanding and experience to the discussion; and Grant is simply the best — and best-known — soccer writer in the U.S."

For more information on the speakers and the program, or to RSVP, visit ucollege.wustl.edu/soccer or call 314-935-6700.

An untold story emerging from cognitive psychology research is that many of us have been going about learning and memorization in all the wrong ways.

That’s a central message of “Make it Stick: The Science of Successful Learning,” a new research-based book that offers students of all ages a clear and compelling primer on the best and worst ways to store and retrieve new knowledge.

Released this month by Harvard University Press/Belknap, the book is co-authored by psychologists Henry L. “Roddy” Roediger III and Mark A. McDaniel, leading experts on human learning and memory at Washington University in St. Louis, along with nonfiction writer and novelist Peter C. Brown. 

Roediger, PhD, the James S. McDonnell Distinguished University Professor in Arts & Sciences, and McDaniel, PhD, professor of psychology in Arts & Sciences, provided the research for the book. Brown interviewed many people whose personal stories are woven into the narrative to illustrate and explain the underlying science. The resulting text is highly accessible for people of all ages who seek to improve their abilities to learn and remember.

Roediger

“Many popular study techniques rely on lots of re-reading and repetitive review in an attempt to force new material into memory, but research shows that gains from this approach are fleeting and often leave student with a false sense of confidence about how well they know a subject,” Roediger said.

“What really matters when it comes to making use of new knowledge is your ability to retrieve it when you need it,” Roediger said. “This book offers concrete tips on study methods that will help you store information in ways that make it more accessible for the long term. 

"One of our main points is that students need to practice retrieving information as a regular part of studying. This way they will have the information at their mental fingertips for use when they need it. Retrieval practice at spaced intervals also slows forgetting. Active retrieval is critical for long-term retention.”

McDaniel

“While much remains to be known about learning and its neural underpinnings, a large body of research has yielded principles and practical strategies that can be put to work immediately, at no cost, and to great effect,” the authors suggest in the book’s preface. “This is a book about what people can do for themselves right now in order to learn better and remember longer.”

Among the findings:

• Recalling new material from memory is more productive for robust learning and memory than rereading.
• Quizzing students on material they have read or that has been presented in class leads to better learning and retention than reviewing that material again.
• Interleaving the practice of two or more problem types (for example, math solutions or types of baseball pitches) is more productive than mastering one before moving to another.
• Asking students to grapple with solving a new kind of problem before teaching them the solution results in better learning of the solution when they are shown it.
• A series of cumulative, low-stakes or no-stakes quizzes over the course of a semester works like compound interest, strengthening retention and updating learning.

These strategies, which require more effort and delay learning, are seldom adopted because learners often mistake the increased effort to be an indication of poor learning, whereas the truth is just the opposite — the increased effort leads to stronger learning and better versatility in recalling and applying the knowledge or skill when it’s needed later.

A Washington University in St. Louis team won the People’s Choice and second place awards at the 2014 Rube Goldberg Machine Contest. The national competition was held at the Center of Science and Industry in Columbus, Ohio, earlier in April.

The contest is named after the late cartoonist Reuben Lucius Goldberg, who died in 1970. For 55 years, the engineer-turned-cartoonist drew machines and contraptions that satirized the technology-infatuated culture of his time. His drawings, using simple gadgets and household items already in use, were incredibly complex, but had an ingenious, logical progression to them.

In the words of the inventor, the machines were a “symbol of man’s capacity for exerting maximum effort to achieve minimal results.”

WUSTL’s Green Machine featured solar panels, a bike, windmills, laundry drying on a clothesline, two gardens — one of which (window-box flowers) won the Best Single Step Award — and lots of backyard wildlife.

“The machine is built mostly of recycled materials,” said Amy Patterson, a junior majoring in biology. “The clothes on the clothesline and the squirrel-garden gnome are from the Trading Post on the South 40. The wind turbine is made of PVC found in a university dumpster, and the heads of the tape-measure flowers are made from insulation found in a dumpster. We use a lot of old packaging materials in our machines, such as empty used soda bottles and used cereal boxes. One of the birds and the material for the curtains came from Goodwill.“

“At the end,” Patterson said, “a squirrel jumps onto the solar-paneled roof, slides into the gutter, pushing down a lever, triggering a mousetrap, pulling out a pin, allowing a weight to drop, zipping the zipper.”

COSI

The Green Machine was designed and built by: (top row, from left) freshman Anish Kanesa-Thasan, junior Amy Patterson, Grace Kuo, a junior majoring in electrical engineering, and (bottom row, from left) Harison Wiesman, a senior physics major and Michelle Heredia, a freshman chemical engineering major. Lydia Zoells (not pictured), a sophmore majoring in English, could not attend the competition.

The machines are judged on story telling, complexity (they must have at least 20 steps), machine flow (can you follow what’s happening), humor, and the use of everyday items for tasks for which they were not intended.

Teams lose points if a team member has to intervene to nudge something that is stuck or if a moveable part “leaves the machine.” 

"This year, we had one perfect run, and one run with two 'touches,' which actually isn't bad," Patterson said.

560 Music Center

Long before John Travolta fumbled her name at the Academy Awards, Idina Menzel was known as the star of “Wicked,” Stephen Schwartz’s musical reimagining of “The Wizard of Oz.” Set years before Dorothy’s arrival, the story centers on Elphaba, a bright young girl of emerald green who grows up to become “the Wicked Witch of the West.”

At 7:30 p.m. Tuesday, April 29, the Washington University in St. Louis Wind Ensemble will present excerpts from “Wicked,” “Phantom of the Opera,” “Star Trek” and more in the Ballroom Theater of the 560 Music Center.

Becker

Continuing the program will be six excerpts from Andrew Lloyd Webber’s “Phantom of the Opera,” as arranged by Paul Murtha. Songs include “Music of the Night,” “The Phantom of the Opera,” “Angel of Music,” “Masquerade,” “Learn to be Lonely” and “All I Ask of You.”

The program will conclude with Brown’s arrangements of the score to the feature film “Star Trek Into Darkness.” Works will include the original television “Theme from Star Trek,” written by Alexander Courage, as well as Michael Giacchino’s “Enterprising Young Men,” “To Boldly Go” and “Ode to Harrison.”

Conducted by Chris Becker, the Wind Ensemble performs both classic and contemporary wind repertoire. Membership is open to undergraduate and graduate students from all WUSTL schools and departments.

The 560 Music Center is located at 560 Trinity Ave., at the intersection with Delmar Boulevard in University City. Milk and cookies will be served after the performance. For more information, call 314-935-5566 or email daniels@wustl.edu.

Whitney Grither, an MD/PhD student at the School of Medicine, accepts her team’s award at the National Cancer Institute (NCI) of the National Institutes of Health (NIH) in March. Also pictured are Tom Stackhouse, associate director of the NCI's technology transfer center, and Rosemarie Truman, founder and CEO of The Center for Advancing Innovation.

The resources required to get discoveries from “bench to bedside” can be astounding. Consequently, many promising innovations never reach their potential. Instead, for a variety of reasons, they stall.

In an effort to reverse this trend, the Avon Foundation for Women, the National Cancer Institute (NCI) and the Center for Advancing Innovation created the Breast Cancer Startup Challenge. The initiative was aimed at bringing 10 promising breast cancer discoveries out of the lab and closer to market to help patients. 

Some 200 teams of graduate students from around the world entered the competition. Forty-six were selected to submit business plans and “elevator pitches” explaining how they would commercially advance any one of these discoveries. A Washington University in St. Louis team was one of the competition’s winners and has been invited to launch a startup based on its plans.

Graduate students Hirak Biswas and Anurag Agarwal in the Division of Biology and Biomedical Sciences and Whitney Grither in the Medical Scientist Training Program won for their proposal to advance a therapeutic breast cancer vaccine developed by NCI researchers and patented in 2011.

“We looked at the competition within the marketplace, as well as other vaccines that have gotten through the FDA regulatory process, to see what we might do to get the product closer to market,” said Grither.

The vaccine is designed to fight cancer by strengthening the body’s immune system to recognize tumor cells as foreign and attack them. It showed promise in preclinical mouse trials, but further research and development is needed before the vaccine is commercially viable and can be used to treat patients. This would be the focus of a startup company. 

Samantha Van Hove

(From left) WUSTL’s Anurag Agarwal, Whitney Grither and Hirak Biswas show their award from the challenge.

The faculty’s “input and expertise in tumor immunology, developing cancer vaccines and corresponding clinical trials were crucial for the team to develop a research and development strategy to develop the technology further,” wrote Biswas in a blog post about the experience.

Evan Kharasch, MD, PhD, vice chancellor for research and the Russell and Mary Shelden Professor of Anesthesiology, praised the students’ accomplishment and said their participation in the challenge “exemplifies Washington University’s initiative on research innovation and entrepreneurship, particularly the role of trainees in advancing promising science into commercialization.”

Sid hastings/WUSTL Photos (2)

Washington University in St. Louis Provost Holden Thorp delivered the 2014 Phi Beta Kappa/Sigma Xi Lecture on April 17 in Simon Hall's May Auditorium.

Thorp gave many examples of why the humanities are essential for a 21st century education, including a clip from a Ken Burns documentary that featured "Star Wars" creator George Lucas, who said: “Science is the ‘how,’ humanities is the ‘why.’”

Jennifer Smith, PhD, dean of the College of Arts & Sciences and associate professor of earth and planetary sciences, introduced Thorp. Vincent Sherry, PhD, the Howard Nemerov Professor in the Humanities and professor of English in Arts & Sciences, moderated the Phi Beta Kappa ceremony.

The Phi Beta Kappa/Sigma Xi Lecture, held April 17 in Simon Hall, was the final Assembly Series program of the semester. Held annually, the lecture is part of the Phi Beta Kappa initiation ceremony. This year, 81 juniors and seniors were inducted into the prestigious society that celebrates excellence in the liberal arts and sciences.

Sid Hastings/WUSTL Photo

Soon Goo Lee and Joseph Jez examining crystallization wells in the cold room of the Jez lab at Washington University in St. Louis. Each well holds a different solution of chemicals and, if they are lucky, one of the solutions will have coaxed a drop of protein hanging from a well lid to crystallize. Once the protein is “frozen” in this way, it can be “photographed” by X-ray diffraction.

If you pull up a soybean or bean plant and shake off the dirt, you might see odd swellings or bumps, like rheumatic finger joints, on its roots. Inside the cool, soil-covered bumps are bacteria that are making nitrogen with the help of an enzyme, something chemical factories can do only with the help of a catalyst and at high temperature and pressure.

The bacteria, typically members of the genus Rhizobia, break the strong triple bond between the nitrogen molecules in the air and repackage the nitrogen atoms in chemical compounds the plant can use. In return, the plant supplies the bacteria with the energy needed to split the nitrogen molecules in the form of sugar.

Legume-Rhizobia partnerships generate more nitrogen for plants than all industrial fertilizers used today, and they provide the right amount of nitrogen at the right time.

By contrast, much of the synthetic fertilizer applied to farm fields is wasted, washing out the soil into waterways or evaporating into the atmosphere in the form of nitrous oxide, where it becomes an environmental and health risk.

Farmers can already buy Rhizobia-rich biofertilizers to increase nodule formation and improve soil quality without synthetic fertilizers. But scientists are beginning to talk about re-engineering crop plants so that, like legumes, they will have on-site nitrogen-fixing systems, either in root nodules or in the plant cells themselves.

To do that, scientists need to understand the biological nitrogen-fixing machinery as thoroughly as a mechanic understands the valves and pistons of a car engine. The difference is that the biological machinery is far too small to be visible to the unaided eye.

Science got one step closer to this goal recently when a team at Washington University in St. Louis worked out the structure of a protein called NolR that acts as a master off-switch for the nodulation process. By building an accurate atomic model of the protein, they were able to “see” exactly how it recognizes and slots itself into genes to prevent  bacteria from embarking on a life as a symbiont. The work was published in the online April 14 edition of PNAS,

Trust but verify
The process of nodulation is so weird that if you read about it in a science fiction book, you’d probably credit the author with a great imagination.

Hari Krishnan

An iron-containing protein not unlike the hemoglobin in our red blood cells colors root nodules. Called leghemoglobin (leg- for “legume”) it ferries oxygen to the respiring bacteria, just as hemoglobin ferries oxygen to our respiring cells. But leghemoglobin holds oxygen much more tightly than hemoglobin and dispenses it more stingily because oxygen can poison the bacterial enzyme that fixes nitrogen.

First, the plant and soil bacteria engage in a molecular dialog to make sure they are compatible partners. The host plant releases a cocktail of chemicals called flavonoids that are perceived by a bacterial protein named NodD, which turns on nod (nodulation) genes. Together the nod genes make a large, complicated molecule called a nod factor.

The nod factor triggers the plant  to make an“infection thread,” or a tube through which the bacteria travel deep into the root, where they are wrapped in membrane the plant has synthesized and sequestered in vesicles within the root cortex cells of a nodule. Their metabolism and even their ability to reproduce are so altered that they are like different organisms, and are called bacteroids rather that bacteria.

“It’s like a pathogen invasion,” said Joseph Jez, PhD, associate professor of biology in Arts & Sciences at Washington University, “but Rhizobium is not a pathogen, it’s a beneficial organism. “

Does it curl or lie flat?
Scientists working on the genetics of nodule formation in the mid-1980s and early ’90s, identified NodD, a master on-switch for nod genes, and NolR, a master off- switch that acts even more broadly, turning off the nod genes, NodD, and other genes needed for life as a symbiont.

“The bacteria has the ability to turn on a bunch of genes for nodulation and the symbiosis, but it needs to keep them turned off as long as it is free-living,” Jez said. “And that’s the role of NolR.”

“It’s like driving a car,” said Soon Goon Lee, PhD, postdoctoral research associate in the Jez lab. “NodD is the gas pedal and NolR is the brake. But we didn’t know how NolR worked until we solved its structure and could understand how it interacted with DNA.”

The job is to figure out how the long, thread-like protein molecule folds on itself to form a snarl of helices and ribbons, and then how the folded molecule fits into and binds with the DNA.

Unfortunately, protein folding is a notoriously hard problem, one as yet beyond the reach of computer calculations. So most protein structures are still solved by the time-intensive process of crystallizing the protein and then barraging the crystal with X-rays to locate the atoms within it.

In 2005, Hari R. Krishnan, PhD, a U.S. Department of Agriculture scientist at the University of Missouri-Columbia, told Jez that he had a NolR clone, or population of cells that all made the NolR protein. Would Jez be interested in trying to crystallize NolR so that its structure could be solved, Krishnan asked.

Jez happily accepted, and, as time permitted, he or another member of the lab would put NolR through a crystallization screen, depositing drops of the protein into little wells filled with different solutions, searching for conditions that would cause the protein to crystallize.

At first, NolR was uncooperative. If they got crystals, the crystals fell apart when they were manipulated or they produced diffraction patterns that couldn’t be interpreted.

In his turn, Lee picked up the project. He decided to start at the beginning again, picking a DNA sequence NolR bound, ordering that snip of DNA, and then trying to crystal a mixture of the protein and DNA together.

This should have been harder than crystallizing the protein alone, but to his surprise it turned out to be easier. The low-resolution data Lee got for the protein/DNA complex made it easier for the scientists to interpret the high-resolution data from the protein alone. 

The protein turned out to have what is called a helix-turn-helix motif commonly found in proteins that bind DNA. The DNA double helix has a major groove and minor groove that run down the double helix like the threads of a screw. Many proteins that bind to DNA do so through the wider major groove.

“The major groove is the one that’s open, and you can fit a protein helix into that groove,” Jez said. “So nature uses this helix-turn-helix domain as a way of positioning helices into the major grooves. The protein is a dimer, so it has two helices that are spaced perfectly to put one into each of two consecutive major grooves.”

To act as a master switch, NolR has to be able to recognize and bind to many different genes. It’s able to do that because each of the genes carries the same sequence of nucleotides, called a consensus sequence, somewhere along its length. In this case, there are two such sequences in consecutive major grooves in all the genes NolR binds.

The scientists are pleased with their progress but it has only made them more eager to crystallize the other protein: the on-switch, NodD. After all, it’s hard to drive a car when you only have a brake but no accelerator.

James Byard/WUSTL Photos

A section of one of the solar arrays installed on the roof of North Campus. Click here for a high-res version.

“These solar projects represent Washington University’s first major investment in renewable energy,” said Phil Valko, director of sustainability. “The investment will pay off by producing enough emission-free energy to power the electrical usage of 53 average U.S. homes, at a savings of $190,000 over 10 years.”

Based on the carbon intensity of grid electricity in Missouri, which is among the highest in the U.S., this will reduce the university's output the equivalent of 385 metric tons of carbon dioxide per year.

“As a university community, we are making great strides in being better stewards of our resources,” said Henry S. Webber, executive vice chancellor for administration. “Last year, we committed to invest $30 million over seven years to reduce our greenhouse gas emissions 22 percent by 2020. This investment in solar energy is among a series of projects that will help us further reduce our carbon footprint and reduce operating costs.”

Most of the solar arrays are being installed on flat roofs, including the Lofts of Washington University (75 kw), North Campus (75 kw), West Campus (75 kw), the Family Learning Center (25 kw), the 560 Music Building (25 kw) and the University City Children’s Center (50 kw).

In addition, Tyson Research Center will have a 50 kw ground-mounted array, and a 4-kw solar carport will be installed over an electric vehicle charging station in front of Brauer Hall on the Danforth Campus. The carport will power the car chargers and be used in battery storage research by the School of Engineering & Applied Science.

Two WUSTL seniors, Rachel Goldstein and Tyler Loucky, were instrumental in helping to identify off-campus sites that would be feasible for photovoltaic rooftop systems. (See sidebar.)

The solar projects were developed as solar leases, with 304 kw developed in partnership with Microgrid Solar and U.S. Bank and 75 kw developed in partnership with Brightergy. The companies own, install and maintain the arrays for the duration of the lease, with minimal up-front cost to the university. Solar rebates from Ameren Missouri were an important component of the project financing.

WUSTL will directly use the solar-generated electricity at each installation site, reducing the amount of energy each building will draw from the grid. The university will pay pre-determined annual lease payments that will be less than the value of the solar electricity generated by the arrays. The percentage of each building’s total electricity usage that will be generated by solar arrays will vary from 1 to 58 percent, depending on the size and energy intensity of each building, with an average of 15 percent across all the sites.

"Many talented individuals at the university contributed their time and expertise to make this project a reality," Valko said. “It simply would not have happened without support from the treasurer’s office, the general counsel’s office, the resource management team, our facilities teams at Danforth and the School of Medicine, the financial planning team, the real estate team and Tyson Research Center.”

For more information about Washington University’s sustainability initiatives, visit sustainability.wustl.edu.

James Byard/WUSTL photos

Cutting the ribbon at the official "roof breaking" event April 23 are (from left) Pratim Biswas, PhD, the Lucy & Stanley Lopata Professor and chair of the Department of Energy, Environmental & Chemical Engineering in the School of Engineering & Applied Science; senior Tyler Loucky; Himadri Pakrasi, PhD, the Myron and Sonya Glassberg/Albert and Blanche Greensfelder Distinguished University Professor and director of I-CARES (International Center for Advanced Renewable Energy and Sustainability); Chancellor Mark S. Wrighton; Phil Valko, director of sustainability; senior Rachel Goldstein; and Henry S. Webber, executive vice chancellor for administration.