21256_Biology Brochure (Page 1) - Department of Biology

The Department OfBIOLOGY

1 Statement from the Chairman2 Department of Biology: A Short History3 Departmental Programs7 Departmental Facilities11 Faculty Program InterestsThe Cover Photos illustrate several model organismsfor ongoing research in the Department.1Image 1. Brienomyrus brachyistius, a weakly electric fish fromWest Africa. (photo by Robert Lewis) (Carlson lab)2Image 2. A male collared lizard showing a novel yellow coloron an Ozark mountain recently colonized after prescribed forestfires. (Templeton lab)Image 3. Photo shows the widespread species Lupinus chamissonis.(Knight lab)3 4Image 4. GUS reporter gene activity (in blue) indicates that themechanosensitive ion channel gene MSL6 is expressed in thevasculature of the Arabidopsis leaf. (Haswell lab)

Statement from the ChairmanThese are very exciting times for biologists.Conceptual and technical advances now comeso rapidly — and create such great potentialfor solving many long-standing problems inbiology, medicine, agriculture, and the environment— that many are already calling the21st century “The Century of Biology.”The Washington University Department ofBiology is in an exceptionally strong positionto capture the tremendous advancementacross and between the subdisciplines of biology:from macromolecular structure, tocell/developmental biology, to ecology, and toevolution. Intellectually, the department drawsits strength from an unusually interactive andcollaborative faculty with a wide range ofinterests at all levels of biological organizationand utilizing many different biologicalsystems. Organizationally, it benefits from itsstrategic location at the intersection of twoadministrative units that are designed to promoteinterdisciplinary interactions in researchand training: as a member of the Division ofNatural Science and Mathematics in Arts &Sciences, Biology is linked to strong physicalscience and math departments; and as amember of the Division of Biology andBiomedical Sciences (DBBS), it is linked tothe basic science departments in one of thecountry’s premier medical schools. New initiativesare reaching out in Engineering as well,e.g. in Imaging Sciences.In recent years productive interactionshave also begun to flourish with the schoolsof Law (patents/biotechnology) and Business(commercial use of biotechnology). Furthermore,the department benefits greatly fromsynergistic interactions that its facultymembers enjoy with colleagues at severaloutstanding off-campus research facilities,including the Donald Danforth Plant ScienceCenter, Monsanto, the Missouri BotanicalGarden, the Tyson Research Center, and theShaw Nature Reserve (all of which are discussedin more detail on the following pages).These interactions ensure our students —from undergraduates to postdoctoral fellows— access to the broadest opportunities fortraining and career development.Our present faculty of 30 includes severalFellows of the AAAS and members of the U.S.National Academy of Sciences. Many of ourfaculty have editorial responsibilities formajor scientific journals, sit on NIH or NSFpeer-review panels, organize national andinternational meetings, and/or have otherleadership positions in the scientific community.One indicator of the breadth of interestsencompassed within the department is thatmany of our faculty members have chosen toaffiliate with eight of the 12 different DBBStraining programs (see http://dbbs.wustl.edu).About 60 graduate students are pursuing PhDdegrees with our faculty while postdoctoralfellows number more than 70.The six contiguous buildings that nowhouse the Biology Department include stateof-the-artfacilities for computing, microscopy,animal and plant growth, a user-friendlylibrary with more than 50,000 boundvolumes and subscriptions to 400 journals,and a Natural Science Learning Center thatserves our more than 400 undergraduatebiology majors. Within the last three yearswe have added eight assistant professorsand one endowed professor jointly with theDepartment of Chemistry, with specializationsin plant science, neurobiology, microbiology,population biology, and ecology. New spaceacquired in 2006 ensures adequate facilitiesto reach our target 34 faculty within three tofive years.In summary, current faculty in the departmentare continuing the long tradition ofexcellence in research and teaching duringthe latter half of the 20th century. The lifesciences at Washington University continueto form a major component of the institution’sintellectual commitment. In this increasinglyinterdisciplinary effort, the Department ofBiology has played and continues to play amajor role.Ralph S. QuatranoSpencer T. Olin Professor and Chairmanwww.biology.wustl.edu/faculty/quatrano.html1

Department of Biology:A Short HistoryThroughout the latter half of the 19th andmuch of the 20th centuries, what we now callthe Department of Biology consisted of separateBotany and Zoology units, both of whichnurtured strong research and teachingprograms.Henry Shaw, founder of the MissouriBotanical Garden in 1859, created the HenryShaw School of Botany at WashingtonUniversity in 1885 and endowed theEngelmann Chair of Botany. Throughout itshistory, the Engelmann Chair has been heldby the director of the Missouri BotanicalGarden and has included such notablebotanists as Fritz Went (who carried out thefirst major studies of the plant growth regulatorauxin) and Edgar Anderson (an evolutionarybiologist who studied hybridization asa mechanism of plant speciation). It is nowheld by Peter H. Raven, a world-renownedchampion of biodiversity. Other distinguishedmembers of the Botany Department haveincluded environmentalist Barry Commonerand microbiologist Howard Gest (one of theearly pioneers on the biochemical pathwaysinvolved in prokaryotic photosynthesis andnitrogen fixation).The Zoology Department also had a distinguishedhistory, particularly after 1935when it was joined by Viktor Hamburger(1900-2001), a former doctoral studentof Hans Spemann in Germany. In 1941Hamburger became chairman, succeedingFrancis O. Schmitt (a leader in the developingfields of molecular neurobiology and biophysics).Hamburger soon hired severaljunior colleagues, all of whom went on torenowned careers as scientific investigators,teachers, and/or administrators. These includedHampton Carson and Harrison Stalker(who collaboratively produced several landmarkstudies in evolutionary biology), H.Burr Steinbach (later chairman of biology atthe University of Chicago, director of theMarine Biological Laboratory in WoodsHole, and president of the Woods HoleOceanographic Institution), Florence Moog(a leader in the emerging field of biochemicalembryology), and Thomas Hall (who becamean eminent administrator and science historian).In the late ’40s Hamburger invited RitaLevi-Montalcini to come from Italy to join hislaboratory as a postdoc. Together with biochemistStanley Cohen (who joined them inthe early ’50s), they initiated the studies thatled to the discovery of nerve growth factor,epidermal growth factor, and the 1986 NobelPrize in Physiology or Medicine for Levi-Montalcini and Cohen. Levi-Montalcini joinedthe faculty in the early ’50s and remained amember until her retirement in the ’80s.In 1969 the Botany and ZoologyDepartments merged to become an integratedBiology Department under the chairmanshipof Johns Hopkins, III. In the 1970s, 1980s,and 1990s, investigators such as NationalAcademy of Sciences (NAS) members JoeVarner, Mary Dell Chilton, Roy Curtiss, III,Barbara Schaal, and Roger Beachy (now directorof the Donald Danforth Plant ScienceCenter in St. Louis) joined the faculty andhelped sustain its long-standing tradition inplant and microbial research. The departmentcurrently boasts four members of the NAS(Beachy, Raven, Schaal, and Suga). Althoughits research tradition is long and strong, thedepartment has always maintained an equallystrong commitment to instruction and trainingof undergraduate and graduate students in abroad range of biological disciplines.In short, the life sciences have alwaysconstituted a major component of the institution’sintellectual and educational commitment.The Department of Biology is proud tobe a major contributor to this increasinglyinterdisciplinary effort.2

DepartmentalProgramsUNDERGRADUATE PROGRAMA biology major provides excellent preparation forstudents interested in careers in medicine, dentistry,biological or biomedical research, biotechnology,bioinformatics, the pharmaceutical industry, agriculture,ecology, conservation, or any of several paramedicalspecialties.Biology is a diverse field in which living systemsare studied in many ways, and at many levels, rangingfrom the molecular and cellular to the population andecosystem levels. Members of the biology faculty arerecognized internationally for their research and bring avariety of strengths and teaching styles into the classroom.Particular areas of strength in the department areplant biology, physiology, molecular genetics, developmentalbiology, and evolutionary biology.RESEARCH OPPORTUNITIESFOR UNDERGRADUATESUndergraduate Alison Goldberger is using a luminometer,which is used to study bacterial enzymaticreactions in the Kranz lab.A wide range of research opportunities isavailable to biology students. Many find theindividual instruction and mentoring receivedwhile doing research to be a centrally importantpart of their undergraduate experience.Since there are more than 350 faculty membersdoing research in biology and biomedicalsciences at Washington University, it is easyto find a project that suits most any particularinterest. Many students complete theirresearch projects at the WashingtonUniversity School ofMedicine, one of thetop-ranked medicalschools in the country.Students often doresearch for credit (Bio200/500, see on nextpage) or as paid work,sometimes as part ofthe federal work-studyprogram. As indicatedon pages 5-6, fundingfor undergraduatesengaging in full-timesummer researchcomes from severalsources, includingHoward HughesMedical Institute, the Children’s DiscoveryInstitute, the WU Office of UndergraduateResearch (OUR), and an NIH training grantin Imaging Sciences. In addition, many researchmentors pay students from research grants forsummer and/or academic year work.The Life Sciences Building contains the NaturalSciences Learning Center, the Biology DepartmentLibrary, and the Jeanette Goldfarb Plant GrowthFacility and Greenhouses.3

T.J. Krall, senior biology major, platesmammalian glial cells to study their rolein circadian rhythms in the Herzog lab.T.J.’s undergraduate research has beensupported by the HHMI summer undergraduateresearch fellowship, theImaging Sciences Training Pathway,and the NSF.BIO 200, BIO 500These academic year courses, called“Introduction to Research” (taken by freshmenand sophomores) and “IndependentWork” (taken by juniors and seniors), allowWashington University students to receiveacademic credit for research carried out in thelab or field-research program of a facultysponsor. Normally, students sign up for threeunits of credit (the equivalent of an averagecourse) and are expected to work 10-12hours per week in the lab plus necessaryreading. Additional information canbe found on the Web at http://www.nslc.wustl.edu/courses/BIO500/bio500/html.Professor Tiffany Knight, right, with senior biologymajor Jocelyn Tsai, discussing the invasive plantLespedeza cuneata. The Knight lab is conductingan experiment that examines the role of nutrientsand interspecific competition in its invasion.LIFE SCIENCESThe Biology Department offers severaloptions for experiences in life sciences.They include Biology 265, Biology 2651, andBiology 2652. Each of these offers a differentopportunity to earn credit for nonclassroomlearning in the life sciences.For BIO 265 a wide variety of activitiesqualify. For example, students might accompanya physician on rounds and prepare apaper on a specific organ system or disease,help create a summer biology curriculum forchildren and report on its effectiveness,intern at a veterinary clinic, etc. Participantsmust arrange to work with a supervisor withwhom they will meet on a regular basis andcommit to at least 140 hours over twosemesters with papers due at the end of thefirst semester and a final report due at theend of the second semester.BIO 2651, MedPrep, is a unique programdesigned specifically for students consideringa career in medicine. The didactic componentconsists of a weekly two-hour lecture on theDanforth Campus.Taught by a physician and member ofthe School of Medicine’s Committee onAdmissions, students are given detailedinformation regarding every step of the medicaleducation process, from applying to medicalschool to becoming a board certifiedphysician. Students learn what doctors likeand don’t like about their chosen specialtiesand the challenges they’ve faced during theirtraining. Topics such as the growing medicalmalpractice crisis, universal health care, andhow physicians cope with death and dyingare also covered. Q&A sessions are also heldwith resident physicians from several specialtiesas well as current medical studentsfrom Washington University.To complement the information presentedin class, students also shadow physicians inthe Emergency and Trauma Center of Barnes-Jewish Hospital three hours every otherweek. Here they see firsthand how physicianstreat both routine medical problemsand life threatening emergencies. Throughthe combination of these experiences studentsget a realistic, uncensored view ofwhat a life in medicine entails. The courseis during the fall and spring semesters only.4

Mark Freeman,Medical SciencesTraining Programstudent, sets up hismultielectrode recordingto the biologicalclock. Mark is simultaneouslypursuingan MD and a PhD inneuroscience.Nicole Herbst,Colgate student forNSF-sponsoredFIBR summer program,and researchassociate AbhaKhandelwal in theQuatrano lab measuringDNA concentration.In BIO 2652 (the PEMRAP Program),students are research associates (RAs) forWashington University faculty membersconducting clinical research studies in theEmergency Department (ED) at St. LouisChildren’s Hospital. Through their efforts asRAs, students interact directly with patientsand families to help gather patient informationfor research projects during shifts in theED. Topic areas of study vary from semesterto semester and include procedural pain andsedation, head injury, fracture healing, asthma,pediatric trauma, and neonatal sepsis.Students also observe evaluations and proceduresduring their shifts. Additionally, PEM-RAP students may elect to participate inadditional shadowing shifts in the ED. Theseallow students to shadow a resident, fellow,or hospitalist through a two-hour period inthe ED. They also give students an opportunityto interact with physicians in a clinicalsetting and to see common and uncommonpediatric emergency issues. This classincludes weekly lectures at the medical centergiven by medical school faculty members ontopics related toemergency medicineand clinicalresearch. Speciallectures are giveneach semesterand include topicssuch as life inmedicine, gettinginto medicalschool, and ethicsof research.GRADUATE PROGRAMSProspective graduate students whowish to work with one of the biologydepartment faculty members shouldapply to the interdepartmental Divisionof Biology and Biomedical SciencesGraduate Program. The Division emphasizesinterdisciplinary approaches andprovides a stimulating cross-departmentalenvironment for students. The 12 predoctoraltraining programs offered by theDivision are:BiochemistryComputational BiologyDevelopmental BiologyEvolution, Ecology, and PopulationBiologyHuman and Statistical GeneticsImmunologyMolecular BiophysicsMolecular Cell BiologyMolecular Genetics and GenomicsMolecular Microbiology and MicrobialPathogenesisNeurosciencePlant BiologyEach program establishes certainprogram-specific requirements forearning the PhD degree, and monitorsthe progress of each of its studentstoward the degree, but the GraduateSchool of Arts & Sciences sets generalizedrequirements and grants the degree.For more detailed information, visithttp://dbbs.wustl.edu.SUMMER RESEARCH OPPORTUNITIESThe Summer Undergraduate ResearchFellowship Program provides support to WUundergraduates for a summer of research,under the direction of members of theDivision of Biology and Biomedical Sciencesand other Washington University and affiliatedfaculty in the natural sciences. The programoriginated with funding from a HowardHughes Medical Institute UndergraduateEducation grant in 1993. Funding for summerwork now comes from several sources5

High school science teachers investigate alternativeenergy and biofuels in a Science Outreach summerprogram. The Center for Science Outreach promotespartnerships between Washington University andK-12 teachers and schools and is supported bythe National Science Foundation, the NationalInstitutes of Health, the Howard Hughes MedicalInstitute, and other donors.including HHMI, the Children’s DiscoveryInstitute, the WU Office of UndergraduateResearch (OUR), and an NIH training grantin Imaging Sciences.Research opportunities are available inbiochemistry, bioorganic chemistry, biophysics,cell biology, computational biology,developmental biology, ecology, earth andplanetary sciences, evolutionary and populationbiology, immunology, molecular genetics,microbiology and microbial pathogenesis,neurosciences, plant science, imaging science,and related topics in laboratories onthe Danforth Campus and at the medicalschool.Applicants identify a faculty mentor anddevelop a brief research proposal. The BiologyDepartment Natural Science LearningCenter Home Page(http://www.nslc.wustl.edu)is an excellent resource for those seeking anappropriate mentor. Follow the links:Research Opportunities, Howard HughesMedical Institute, Imaging Sciences Pathway,How to Find a Mentor. (Further assistance, ifneeded, is available from the Bio200/Bio500faculty advisor.)This program provides a stipend for aminimum of 10 weeks of full-time effort.Some fellows live as a group on the DanforthCampus in a specialdorm that houses studentsdoing researchin any area of study.The WU OUR providesa partial housingsubsidy to studentschoosing thisoption. Living in acommunity with othersengaged inresearch enhances theexperience. The OURalso organizes a numberof social eventsand activities for studentsengaged insummer research.Fellows may notenroll in courses oraccept other employmentduring the 10-week period.Students with prior research experienceand those new to research are both encouragedto apply. An important goal of this programis to encourage women and membersof groups that are presently underrepresentedin biomedical research (includingAmerican Indians, African Americans,Hispanics, Native Alaskans, Native PacificIslanders, or persons with disabilities) toparticipate in research in order to evaluatetheir interest in a career in biomedicalscience.SUMMER RESEARCH SPONSORED BYINDIVIDUAL LABSMany students who start in a lab as workstudy students or by doing Bio 500 are providedfinancial assistance by their facultymentors to continue their work during thesummer. Sometimes they do technical work,but often they do research. The NationalInstitutes of Health and the National ScienceFoundation will often supplement facultyresearch grants to provide summer supportfor undergraduate research students.SCIENCE OUTREACH PROGRAMThe mission of the Washington UniversityScience Outreach program is to link thephysical and intellectual resources of theUniversity to community schools, with thegoal of enhancing the teaching of investigativescience in K-12 classrooms. To that end,the Outreach Office provides a number ofdifferent programs that are designed to promoteaccurate, up-to-date, and enjoyableinquiry-based science instruction at thepre-college level.These programs include developmentand implementation of inquiry-basedcurricula, academic-year and summerprofessional-development courses for K-12teachers, facilitation of teacher-networkingmeetings, and operation of a clearinghousedesigned to deal with the many requests forinformation, expendable supplies, loanerequipment, and classroom instructionalsupport. For more information, visithttp://www.so.wustl.edu.6

DepartmentalFacilitiesThe Department of Biology is housed in seven buildingswith more than 135,000 square feet of space. Support servicesinclude administration, facility management, accounting,and stockroom and purchasing services. In addition, full-timestaff members provide the faculty with support for variousaspects of undergraduate instruction.The Jeanette Goldfarb Plant GrowthFacility is a 10,000-square-foot facility consistingof greenhouses, growth chambers,and a plant tissue culture lab. The fivegreenhouses are controlled by an Argus integratedcontrol system and provide a totalplant-growth area of 5,200 square feet. Thegrowth chamber facility houses 22 reach-inand walk-in chambers. The Plant GrowthFacility is managed and maintained by agreenhouse manager with help from staffand students.The Integrated Macromolecular Facility isprofessionally staffed and has a variety ofsophisticated machines, including a phosphoimagerand real-time PCR machine. As partof the Integrated Macromolecular Facility, theDNA sequencing core facility has two automatedmachines for high-throughput DNAanalysis. The first, an ABI 3130x1, is capableof running unattended with a throughputof 350-400 sequencing reactions per day.The second, an MJR BaseStation, is used primarilyfor DNA fragment analysis and canaccurately size up to 10,000 fragments in an8-hour shift.The Biology Computing Facility is staffedby two full-time computer specialists andprovides computing support for all officesand research labs. A high-speed networklinks more than 300 computers throughoutthe department and provides access to theinternet, electronic mail, file sharing, andnetwork printing. In addition, two centralizedProfessor Ken Olsen points out the pattern offloral development in a strain of invasive weedyrice to undergraduate summer scholar Erica Colein the Biology Department greenhouses.labs provide several PC, Macintosh, and Unixworkstations, plus supplemental equipmentincluding flatbed scanners, slide scanners,color printers, and a poster maker. For moreinformation, visit http://www.biology.wustl.edu/biocomputing/index.php.The Natural Sciences Learning Center(NSLC) is designed as a “home base” forundergraduate biology students. It is locatedon the ground floor of the Life SciencesBuilding, close to the library and other teachinglabs. It includes a student lounge, a computerclassroom, a conference room, eighttutorial rooms, and the office of the Bio-Computing Coordinator. The NSLC alsoserves as a centralized source of informationon careers, internships, and research andgraduate programs of potential interest tobiology majors. For more information, visithttp://nslc.wustl.edu.The departmental Microscope Facility ismanaged by a professional microscopist and7

species, the facility also accommodatesbehavioral research and studies that involvehazardous agents. The facility employs threefull-time staff members, two of whom arecertified by the American Association ofLaboratory Animal Science. The staff ismanaged by a laboratory animal technologistand works closely with the Division ofComparative Medicine veterinarians. Its animalcare program complies fully with theAnimal Welfare Act and Public HealthService Policy on Humane Care and Use ofLaboratory Animals.Mamiko Isaji, PhD, Debbie Frank, PhD, and MikeVeith, MS, study images using the confocal Leicalaser scanning microscope. This microscopesystem allows for detection and localizationof fluorescing molecules.houses a transmission and a scanning electronmicroscope, a Leica laser scanning confocalmicroscope, a computer-controlled optical-sectioningmicroscope, a Zeiss Axiomatfor phase and video microscopy, and a photon-countingimaging system for real-timemonitoring of bioluminescence or fluorescencein living single cells.The Biology Library contains books andjournals relating to many different areas ofbiology. Library staff are eager to assist students,researchers, and visitors. The twofloorfacility houses study space, internetworkstations, printers, photocopiers, andvideocassette players. Extensive electronicresources are also available from the library’sweb site. For more information, visithttp://library.wustl.edu/units/biology/.Tyson Research Center is a 2,000-acre fieldstation owned by Washington University andlocated about 20 miles southwest of theDanforth Campus. Tyson consists of rollinghills of oak-hickory forest, as well as severalsmall streams, ponds, old fields, and glades;the scenic Meramec River borders Tyson tothe north. In addition, several other naturalareas surround Tyson, providing a bufferfrom the ever-encroaching suburbs.Tyson provides a hub for a growingresearch program in environmental biologybased in the Department of Biology atWashington University. During thespring/summer field season, a vibrant communityof researchers, including WashingtonChris Shaffer, director of the DNA SequencingFacility, and DBBS graduate student Taina Pricereview data collected with the ABI 3130sequencing machine.The Danforth Animal Facility is an 8,000-square-foot, AAALAC-accredited animal careunit conveniently housed near all of thedepartment’s buildings, and which functionsas a centralized resource for the DanforthCampus. In addition to animal holding areasfor conventional and exotic laboratory8

University faculty, postdoctoral fellows,and graduate and undergraduate students, aswell as researchers from nearby institutions,use Tyson to address some of the mostimportant basic and applied questions inenvironmental biology at the genetic, population,community, landscape, and ecosystemscales. A laboratory and office complex areavailable onsite, and financial and logisticalsupport is available for those wishing to initiateprojects at Tyson. Researchers fromEarth and Planetary Sciences, Physics,Environmental Engineering, Art, Anthropology,the medical school, and severalother departments also use Tyson for variousresearch and teaching activities. In addition,several Washington University courses useTyson for field trips and outdoor laboratories.Tyson Research Center provides opportunities fora wide range of biological studies from genes toecosystems.DONALD DANFORTH PLANTSCIENCE CENTERThe Donald Danforth Plant Science Center (DDPSC)was founded in 1998 as the product of a unique andinnovative partnership among Washington University,the University of Missouri-Columbia, the Universityof Illinois-Urbana/Champaign, Purdue University,the Missouri Botanical Garden, and MonsantoCompany. It is located about six miles from theWashington University campus.Led by world-renowned plant scientist RogerN. Beachy, the mission of the not-for-profitDanforth Center is to increase understanding ofbasic plant biology; to facilitate the rapid developmentand commercialization of promising technologiesand products; to apply new knowledgeof this sort for the benefit of human nutritionand health and to improve the sustainability ofagriculture worldwide; and to contribute to theeducation and training of graduate and postdoctoralstudents, scientists, and technicians from aroundthe world.The Danforth Center’s research facility openedin November 2001 and will ultimately house upto 250 scientists. For more information, visithttp://www.danforthcenter.org/.The terra cotta and glasscladDanforth Center withgreenhouses attached.9

MISSOURI BOTANICAL GARDENFounded in 1859, the Missouri BotanicalGarden is the oldest botanical garden in thecountry and a National Historic Landmark.Its 79 acres of gorgeous landscaping, displaygardens, and historic structures include theClimatron ® tropical rain forest conservatory;Seiwaen, one of the largest authenticJapanese strolling gardens in the UnitedStates; and Tower Grove House, founderHenry Shaw’s original 19th-century estatehome. The Garden is justifiably one ofSt. Louis’s most famous attractions.More than just a pleasant place to visit,the Missouri Botanical Garden is also a centerfor science and conservation, education,and horticultural display, and is rankedamong the top such institutions in the world.With a mission “to discover and shareknowledge about plants and their environment,in order to preserve and enrich life,”the Garden employs 45 PhD scientists whowork in more than 36 countries in a race torecord and preserve biodiversity.The Garden also offers a top botanicalresearch library with an exquisite rare bookcollection; one of the world’s largest herbariumswith 5.8 million mounted plantspecimens; and an online botanicaldatabase that serves researchersaround the globe.The Garden’s president, Peter H.Raven, is considered one of the leadingbotanists in the world. He is aleading advocate for sustainability,conservation, and the preservation ofbiodiversity and has written dozensof books and articles on the plantkingdom. During his 36-year tenure,Raven has proven as adept infundraising and management of theGarden as he has been as a scientist.The Garden’s family of attractions encompassesthree additional St. Louis-areacampuses:Established by the Missouri Botanical Gardenin 1925, the 2,400-acre Shaw NatureReserve is located in Gray Summit, 35 mileswest of St. Louis. Fourteen miles of hikingtrails draw nature and outdoor enthusiasts inall four seasons to the prairie, glade, andwetland ecosystems, and to admire thenative wildflowers.A division of the Garden since 2001, theSophia M. Sachs Butterfly House is locatedin Faust Park, west St. Louis County. Itsglasshouse conservatory features thousandsof colorful tropical butterflies in free flight,as well as educational exhibits. Outdoorhabitats show off native species.The EarthWays Center in midtown St. Louisis a Victorian house renovated to demonstrateenergy efficient systems, recycledproducts, and sustainable lifestyle choices.Tours and school groups may visit the houseone weekend each month or by appointment.The drum bridge (taiko bashi) inthe Japanese Garden at the MissouriBotanical Garden.10

Faculty Program InterestsGraduate Studies: Division of Biology & Biomedical SciencesGarland Allen: Evolution, Ecology andPopulation BiologyRoger Beachy: Molecular Cell Biology,Molecular Microbiology and MicrobialPathogenesisYehuda Ben-Shahar: Molecular Geneticsand Genomics; Molecular Cell Biology;Neuroscience; Evolution, Ecology andPopulation BiologyRobert Blankenship: Biochemistry,Molecular Biophysics, Plant BiologyBruce Carlson: Neuroscience; ComputationalBiology; Evolution, Ecology and PopulationBiologyDouglas Chalker: Molecular Genetics andGenomics, Molecular Microbiology andMicrobial Pathogenesis, DevelopmentalBiologyJon Chase: Evolution, Ecology andPopulation BiologyEllen Damschen: Evolution, Ecology andPopulation BiologyRam Dixit: Molecular Cell Biology, PlantBiologyIan Duncan: Developmental Biology,Molecular Genetics and GenomicsSarah Elgin: Molecular Genetics andGenomics, Developmental BiologyUrsula Goodenough: Plant Biology,Developmental BiologyElizabeth Haswell: Plant Biology; MolecularGenetics and GenomicsErik Herzog: Neuroscience; MolecularCell Biology; Developmental BiologyDavid Ho: Plant Biology; MolecularGenetics and GenomicsTiffany M. Knight: Evolution, Ecology andPopulation Biology; Plant BiologyRobert Kranz: Plant Biology, MolecularMicrobiology and Microbial PathogenesisBarbara Kunkel: Plant Biology, MolecularMicrobiology and Microbial Pathogenesis,Molecular Genetics and GenomicsAllan Larson: Evolution, Ecology andPopulation Biology; Computational Biology;Molecular Genetics and GenomicsPetra Levin: Molecular Microbiology andMicrobial Pathogenesis, Molecular CellBiology, Molecular Genetics and GenomicsKathryn Miller: Developmental Biology,Molecular Cell BiologyKenneth M. Olsen: Evolution, Ecology andPopulation Biology, Plant BiologyJohn Orrock: Evolution, Ecology andPopulation BiologyPhillip Osdoby: Developmental Biology,Molecular Cell BiologyHimadri Pakrasi: Plant Biology,Biochemistry, Computational BiologyCraig Pikaard: Plant Biology, MolecularGenetics and Genomics, BiochemistryRalph Quatrano: Plant Biology,Developmental Biology, Molecular CellBiologyPeter Raven: Evolution, Ecology andPopulation Biology; Plant BiologyEric Richards: Plant Biology, MolecularGenetics and GenomicsBarbara Schaal: Evolution, Ecology andPopulation Biology; Plant BiologyPaul Stein: NeurosciencesAlan Templeton: Evolution, Ecology andPopulation Biology; Computational Biology;Molecular Genetics and Genomics; Humanand Statistical GeneticsRobert Thach: Molecular Genetics andGenomics, Molecular Cell Biology,Biochemistry11

Garland E. AllenProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONProfessor Garland E. Allen’s research interestsare in the area of history and philosophyof biology — particularly genetics, embryology,and evolution — and their interrelationshipsbetween 1880 and 1950. This workfocuses particularly in the early developmentof the Mendelian-chromosome theory as formulatedin the work of T.H. Morgan and hisgroup at Columbia University and later at theCalifornia Institute of Technology. Growingout of this work have been a series of studiesof the scientific, economic, and social historyof “eugenics” (defined in the early partof the 20th century as “the science of humanimprovement through better breeding”). Thehistory of eugenics provides a number ofinsights into the interrelationships betweenscience and its social context, and raisesmany issues of ethical, legal, and socialimportance that are surfacing today in relationto modern genomics and the HumanGenome Project.SELECTED PUBLICATIONS“A Century of Evo-Devo: The Dialectics ofAnalysis and Synthesis in Twentieth-CenturyLife Science” in Evo-Devo, Past and Present.Eds. Jane Maienschein and ManfredLaubichler. Cambridge, MA: MIT Press(2007). 123-167.“Mechanism, Vitalism and Organicism in LateNineteenth and Twentieth-Century Biology:The Importance of Historical Context,” Studiesin the History and Philosophy of Biologicaland Biomedical Sciences 36 (2005). 261-283.“Heredity, Development and Evolution at theCarnegie Institution of Washington,” in TheDepartment of Embryology at the CarnegieInstitution of Washington, A History. Eds.Jane Maienschein, Marie Glitz and Garland E.Allen. Cambridge, UK: Cambridge UniversityPress (2005). 145-171.“Mendelian Genetics and Postgenomics: TheLegacy for Today,” in Ernst Mayr CentennialVolume, Ludus Vitalis XII. Ed. Francisco J.Ayala. (N. 21, Summer 2004). 213-236.“The Classical Gene: Its Nature and ItsLegacy” in Mutating Concepts, EvolvingDisciplines: Genetics, Medicine and Society.Eds. Lisa S. Parker and Rachel Ankeny.Dordrecht, Netherlands: Kluwer AcademicPublishers. (2003). 11-41.Thomas Hunt Morgan: The Man and HisScience. Princeton: Princeton University Press.(1978).12Logo from the program of the Third InternationalCongress of Eugenics, held at the AmericanMuseum of Natural History in New York, August1932. Eugenics was viewed as a tree drawingfrom many disciplines, especially genetics,anthropology, and statistics, for its sources.“Is A New Eugenics Afoot?” Science, 294 (5October, 2001). 59-61.

Roger N. BeachyProfessor of Biology, President & Director,Donald Danforth Plant Science Centerwww.danforthcenter.orgRESEARCH DESCRIPTIONResearch in the Beachy laboratory is directedto understanding how plant cells and tissuesrespond to infection by viruses, includingidentifying how virus replication complexesare formed and how infection moves fromcell to cell. A related project is to define therole of transcription factors and promotersequences that control gene expression of apararetrovirus. Knowledge from these studiesis used to develop mechanisms to limit virusreplication and to develop a chemically controlledgene switching system to regulateexpression of transgenes in plants.Accumulation of the tobacco mosaic virus 30 kDa movementprotein in BY-2 protoplasts following infection. Fluorescencemicroscopy was used to visualize MP:GFP fusion protein inassociation with microtubules and ER.SELECTED PUBLICATIONSBazzini, A., Asurmendi, S., Hopp, H.E. andBeachy, R.E. (2006). TMV and PVX coat proteinsconfer heterologous interference to PVXand TMV infection, respectively. Journal ofGeneral Virology, Vol. 87: 1005-1012.Bazzini, A., Hopp, H.E., Beachy, R.N. andAsurmendi, S. (2006). Posttranscriptionalgene silencing does not play a significantrole in PVX coat protein mediated resistance.Phytopathology, Vol. 96: 1175-1178.Dai, S., Zhang, Z., Bick, J. and Beachy. R.N.(2006). Essential role of box II cis elementand cognate host factors in regulating thepromoter of rice tungro bacilliform virus.Journal of General Virology, Vol. 87: 715-722.Fujiki, M., Kawakami, S., Kim, R.W. andBeachy, R.N. (2006). Domains of Tobaccomosaic virus movement protein essential forits membrane association. Journal of GeneralVirology, Vol. 87: 2699-2707.Kouassi, N.K., Chen, L., Siré, C., Bangratz-Reyser, M., Beachy, R.N., Fauquet, C.M. andBrugidou, C. (2006). Expression of rice yellowmottle virus coat protein enhances virusinfection in transgenic plants. Arch Virology,151: 2111-2122.Petruccelli S., Otegui, M., Lareu, F., Tran Dinh,O., Fitchette, A.-C., Circosta, A., Rumbo, M.,Bardor, M., Carcomo, R., Gomord, V. andBeachy, R.N. (2006). A KDEL-tagged monoclonalantibody is efficiently retained in theendoplasmic reticulum in leaves, but is bothpartially secreted and sorted to protein storagevacuoles in sees. Plant Biotechnology Journal,Vol. 4: 511-517.13

Yehuda Ben-ShaharAssistant Professor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONAnimal behavior is mediated by their nervoussystem according to information encoded intheir genomes. The Ben-Shahar lab uses avariety of behavioral, genetic, genomic, biochemical,and molecular techniques to decipherthe genetic architecture that drives specificbehaviors. We use the powerful model of thefruit fly, Drosophila melanogaster, to understandhow the function of specific genes indistinct neuronal circuits gives rise to behaviorssuch as feeding and mating. We are alsointerested in taking advantage of the recentsequencing of 12 genomes of species from theDrosophila genus, which now allows us to askquestions about the evolution of behavior.Currently the lab is focusing on severalprojects:• The role of ligand-gated ion channels asneuronal modulators and sensory transducers• The role of neuropeptides and their receptorsin modulating behavior• The role of divalent cation transportersin feeding induction and food selection• The evolution of behaviorally relatedgenomic modules in the genus Drosophila.RECENT PUBLICATIONSBen-Shahar, Y., Nannapaneni, K.,Casavant, T.L., Scheetz, T.E. and Welsh, M.J.(2007). Eukaryotic operon-like transcriptionof functionally related genes in Drosophila.Proceedings of the National Academy ofSciences, 104: 222–227.Whitfield, C.W., Ben-Shahar, Y., Brillet, C.,Leoncini, I., LeConte, Y., Rodriguez-Zas, S.and Robinson, G.E. (2006). Genomic dissectionof behavioral maturation in the honeybee. Proceedings of the National Academy ofSciences, 103: 16068–16075.Ben-Shahar, Y., Dudek, N. and Robinson, G.E.(2004). Malvolio, manganese, and division oflabor in honey bee colonies: deconstructing acomplex phenotype. Journal of ExperimentalBiology, 207: 3281-3288. (Cover)Ben-Shahar, Y., Leung, H-T., Pak, W.L.,Sokolowski, M.B. and Robinson, G.E.(2002). Division of labor in honey beecolonies is influenced by cGMP-dependentchanges in phototaxis. Journal ofExperimental Biology, 206: 2507-2515.Robinson, G.E. and Ben-Shahar, Y. (2002).Social behavior and comparative genomics:new genes or new gene regulation? GenesBrain and Behavior, 1:197-203.Ben-Shahar, Y., Robichon, A., Sokolowski,M.B. and Robinson, G.E. (2002). Influenceof gene action across different time scales onbehavior. Science, 296: 741-744.14Larval GFP expression driven by the promoter of lounge lizard, amember of the Degenerin/ENaC family of cation channels. Dottedline represents the outline of the larval head. Note the expressionpattern in two distinct classes of sensory neurons: onemechanosensitive multi-dendritic neurons (md) and two externalchemosensory neurons (es) projecting to the terminal tasteorgan (TO).

Robert E. BlankenshipLucille P. Markey Distinguished Professor in Arts &Sciences, Professor of Biology and Chemistrywww.biology.wustl.edu/facultyElectron microscopictomaogramof a dividing photosyntheticbacterium.RESEARCH DESCRIPTIONOur research program is primarily concernedwith elucidating the mechanism of the energy-storingreactions in photosynthetic organisms,as well as understanding the originand early evolution of photosynthesis.The chemical reactions leading to long-termenergy storage in photosynthetic systemstake place within the membrane-bound reactioncenter complex and an associated groupof proteins that make up an electron transportchain. One of the central goals of ourresearch is to identify the molecular parametersresponsible for the fact that essentiallyevery photon absorbed by the system leadsto stable products. To this end, we do a varietyof kinetic, thermodynamic, and structuralmeasurements on antenna complexes, reactioncenters, electron transport proteins, andisolated pigments, using a number of techniques,including ultrafast laser flash photolysisand UV-VIS, fluorescence and electronspin resonance spectroscopies, as well asbiochemical and molecular biologicalanalysis.The appearance of photosynthesisand other metabolicprocesses such as nitrogenfixation had profound effectson the evolution of advancedlife on Earth. Our analysis ofwhole bacterial genomes hasrevealed that these metabolicprocesses have complex evolutionaryhistories, includingsubstantial horizontal genetransfer. We have also used acombination of genomics,molecular evolution techniques,and biochemical analysis to identifyand characterize previously unknownenzyme complexes with novel activities.RECENT PUBLICATIONSvan de Meene, A.M.L., Olson, T.L., Collins,A.M. and Blankenship, R.E. (2007). InitialCharacterization of the PhotosyntheticApparatus of “Candidatus Chlorothrixhalophila”: A Filamentous, AnoxygenicPhotoautotroph. Journal of Bacteriology, 189:4196-4203.Kiang, N., Siefert, J., Govindjee, Blankenship,R.E. (2007). Spectral signatures of photosynthesis.I. Review of earth organisms.Astrobiology, 7: 222-251.Swingley, W.D., Gholba, S., Mastrian, S.D.,Matthies, H.J., Hao, J., Ramos, H., Acharya,C.R., Conrad, A.L., Taylor, H.L., Dejesa, L.C.,Shah, M.K., O’Huallachain, M.E., Lince, M.T.,Blankenship, R.E., Beatty, J.T. andTouchman, J.W. (2007). The CompleteGenome Sequence of Roseobacter denitrificansReveals a Mixotrophic Rather thanPhotosynthetic Metabolism. Journal ofBacteriology,189: 683-690.Sadekar, S., Raymond, J., and Blankenship,R.E. (2006). Conservation of distantly relatedmembrane proteins: photosynthetic reactioncenters share a common structural core.Molecular Biology and Evolution, 23: 2001-2007.Melkozernov, A.N., Barber, J. andBlankenship, R.E. (2006). Light-harvestingin photosystem I supercomplexes.Biochemistry, 45: 331-345.15

Bruce A. CarlsonAssistant Professor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONOne of the primary functions of nervous systemsis to extract information from the outsideworld and use that information to guideand coordinate behavior. When studying aparticular nervous system, it is thereforeimportant to consider the ecological contextand evolutionary forces that have shapedthat animal’s natural behavior. The Carlsonlab combines electrophysiology, neuroanatomy,computational modeling, and behavioralanalysis to study information processing inthe electrosensory systems of weakly electricfish from Africa and South America. Theseunique creatures are well suited to establishingdirect links between the physiology ofindividual neurons and quantitative characteristicsof natural behaviors, and are thereforean attractive model system for addressingbasic questions about information processingby sensory systems. How do theactivity patterns of peripheral sensory neuronsrepresent information about the outsideworld? How do central sensory neuronsextract biologically relevant information fromthese patterns of activity? How do centralsensory neurons integrate information frommultiple sources? By focusing on specificresearch topics related to these broad questions,we are beginning to elucidate theimpressive computational power of nervoussystems.RECENT PUBLICATIONSCarlson, B.A. and Kawasaki, M. (2006).Stimulus selectivity is enhanced by voltagedependentconductances in combination-sensitiveneurons. Journal of Neurophysiology,96: 3362-3377.Carlson, B.A. and Kawasaki, M. (2006).Ambiguous encoding of stimuli by primarysensory afferents causes a lack of independencein the perception of multiple stimulusattributes. Journal of Neuroscience, 26: 9173-9183.Arnegard, M.E. and Carlson, B.A. (2005).Electric organ discharge patterns duringgroup hunting by a mormyrid fish.Proceedings of the Royal Society B:Biological Sciences, 272: 1305-1314.Carlson, B.A. and Hopkins, C.D. (2004).Central control of electric signaling behaviorin the mormyrid Brienomyrus brachyistius:segregation of behavior-specific inputs andthe role of modifiable recurrent inhibition.Journal of Experimental Biology, 207: 1073-1084.Carlson, B.A. and Kawasaki, M. (2004).Nonlinear response properties of combination-sensitiveelectrosensory neurons in themidbrain of Gymnarchus niloticus. Journal ofNeuroscience, 24: 8039-8048.Carlson, B.A. and Hopkins, C.D. (2004).Stereotyped temporal patterns in electricalcommunication. Animal Behaviour, 68: 867-878.16Brienomyrus brachyistius, a weakly electric fishfrom West Africa (photo by Robert Lewis).Carlson, B.A. (2002). Electric signalingbehavior and the mechanisms of electricorgan discharge production in mormyrid fish.Journal of Physiology—Paris, 96: 403-417.

Douglas L. ChalkerAssistant Professor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONOnly a fraction of the genome is made up ofactual genes. What functions are associatedwith non-genic DNA, and how do cells dealwith “extra” DNA? All cells must recognizeboth coding and non-coding DNA and organizeit for proper gene regulation. Recent discoverieshave revealed that RNA moleculescan dramatically influence the activity of thegenome, guided by their sequence complementarityto specific regions. In the ciliateTetrahymena thermophila, complementary,small RNAs direct massive remodeling of thedeveloping somatic genome, a phenomenonmechanistically related to RNA interference(RNAi). These small RNA molecules targetspecific chromatin modifications to thehomologous regions, and the DNArearrangement machinery recognizes themodified chromatin state and eliminates thetargeted DNA segment. Our lab aims tounderstand the regulation of this massivegenome reorganization using a combinationof genetic, molecular, and cellular biologyapproaches. These studies will certainly providefundamental insight into RNAi-relatedmechanisms that direct chromatin modificationsthat are critical for transcriptional genesilencing and heterochromatin formation ineukaryotes. Underlying this work is a goal tounderstand how RNA molecules can communicategenetic information between theparental and developing genomes, which hasgreat potential to reveal novel roles for RNAin epigenetic programming. Additionally, webelieve many of the DNA segments targetedfor elimination are important for germ linechromosome structure, and thus understandinghow the cell specifically recognizes thesesequences will contribute general knowledgeof mechanisms ensuring chromosome stabilitythat are essential to prevent aberrantrearrangements.SELECTED PUBLICATIONSYao, M-C., Yao, C-H., Halasz, L.M., Fuller, P.,Rexer, C.H., Wang, S.H., Jain, R., Coyne, R.S.and Chalker, D.L. (2007). Identification ofnovel chromatin-associated proteins involvedin programmed genome rearrangements inTetrahymena. Journal of Cell Science, 120(9):1978-1989.Meyer, E. and Chalker, D. L. (2007).Epigenetics of Ciliates. In Epigenetics, Eds C.D. Allis, T. Jenuwein, D. Reinberg and M.-L.A. E. Caparros. Cold Spring Harbor: ColdSpring Harbor Press. 127-150.Kowalczyk, Christina A., Anderson, AlissaM., Arce-Larreta, Maria, Chalker, Douglas L.(2006). The germ line limited M element ofTetrahymena is targeted for elimination fromthe somatic genome by a homology-dependentmechanism. Nucleic Acids Research,34(20): 5778-5789.Malone, C.D., Anderson, A.M., Motl, J.A.,Genome rearrangements involve dynamic nuclear Rexer, C.H. and Chalker, D.L. (2005).reorganization. Shown here is a Tetrahymena Germline transcripts are processed by acell during genome reorganization. The DNA Dicer-like protein that is essential for developmentallyprogrammed genome rearrange-rearrangement protein, Pdd1p, fused to greenfluorescent protein, is localized on the chromatinments of Tetrahymena thermophila.within the large, developing somatic nuclei.Molecular Cell Biology, 25(20): 9151-64.Pdd1p-GFP localizes in punctate foci in whichDNA elimination occurs. Using GFP fusions tovisualize DNA rearrangement in live cells revealsthe striking dynamics of this process. 17

Jon M. ChaseAssociate Professor of BiologyDirector, Tyson Research Centerwww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONDo rules govern ecology, or are we simplynatural historians gathering special cases?Jon Chase’s research interests are broad butgenerally focus on the rules (or lack thereof)underlying the diversity, distribution, andabundance of animal and plant species fromthe population/community/ecosystem perspective.He is particularly interested in thepatterns and processes that develop at theinterface between local and regional spatialscales. To approach these questions, he combinesmathematical theory, observations andstatistical approaches, rigorous experimentationin both the field and lab, and a knowledgeof natural history.SELECTED PUBLICATIONSChase, J. M. (2005). Towards a really unifiedtheory for metacommunities. FunctionalEcology, 19: 182-186.Johnson, P.T.J. and Chase, J.M. (2004).Parasites in the food web: linking amphibianmalformations and aquatic eutrophication.Ecology Letters, 7: 521-526.Chase, J. M. (2003). Community assembly:when does history matter? Oecologia, 136:489-498.Chase, J. M. and Leibold, M.A. (2003).Ecological Niches: Linking Classical andContemporary Approaches. University ofChicago Press, Chicago, IL.Chase, J. M. and Leibold, M.A. (2002).Spatial scale dictates the productivity-diversityrelationship. Nature, 415: 427-430.Four mesocosms at the Tyson Research Centerwith exactly the same environmental conditions.Note the difference in the macrophyte speciesdominating each community (amphibians andinvertebrates also vary considerably). The onlydifference among the mesocosms is the order inwhich species entered the communities, and thesemultiple community states have maintainedthemselves for more than five years.18

Ellen I. DamschenAssistant Professor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONHow do the environment and spatial processesinteract to determine community composition?How are humans changing these interactionsand what does this mean for globalbiodiversity? The Damschen lab asks howand when space matters for the diversity andcomposition of communities, especially underthe ever increasing impact of humans on theglobe. Our research lies at the intersection ofproviding empirical tests of ecological theoryand providing scientific information to conservationmanagers. Current projects in thelab include:1) How corridors and edge effects affectplant communities2) Using species traits to predict landscaperesponses3) How climate change affects edaphicendemic plants4) How connectivity varies acrossecosystemsOur study sites include the Savannah RiverSite near Aiken, South Carolina; theKlamath-Siskiyou Mountains in southwesternOregon; the kelp forests of the SantaBarbara Channel; Tyson Research Center nearSt. Louis, Missouri; and the Ozark glades.RECENT PUBLICATIONSDamschen, E.I., Haddad, N.M., Orrock, J.L.,Tewksbury, J.J. and Levey, D.J. (2006).Corridors increase plant species richness atlarge scales. Science, 313(5791): 1284-1286.Orrock, J.L., Levey, D.J., Danielson, B.J. andDamschen, E.I. (2006). Seed predation, notseed dispersal, explains the landscape-levelabundance of an early-successional plant.Journal of Ecology, 94: 838-845.Damschen, E.I., Rosenfeld, K.M., Wyer, M.,Murphy-Medley, D., Wentworth, T.R., andHaddad, N.M. (2005). Visibility matters:Student knowledge of women’s contributionsto ecology. Frontiers in Ecology and theEnvironment, 3(4): 212-219.Orrock, J.L. and Damschen, E.I. (2005).Corridors cause differential seed predation.Ecological Applications, 15(3): 793-798.Tewksbury, J.J., Levey, D.J., Haddad, N.M.,Sargent, S., Orrock, J.L., Weldon, A.,Danielson, B.J., Brinkerhoff, J., Damschen,E.I. and Townsend, P. (2002). Corridorsaffect plants, animals, and their interactionsin fragmented landscapes. Proceedings ofthe National Academy of Sciences USA, 99(20): 12923-12926. Awarded the 2002Outstanding Paper in Landscape Ecology,International Association of LandscapeEcology.Clockwise from the left: The Klamath-Siskiyoumountains near Selma, Oregon; a kelp forest inthe Santa Barbara Channel; and The CorridorProject at the Savannah River Site near Aiken,South Carolina. Photo Credits: PISCO and E.Damschen19

Ram DixitAssistant Professor of Biologywww.biology.wustl.edu/faculty20RESEARCH DESCRIPTIONCell wall-encased plant cells are like bricksthat build plant architecture. However, these“bricks” can transform their shape inresponse to developmental and environmentalcues and thereby dynamically regulateplant morphology. The Dixit lab is interestedin understanding themechanisms that regulateplant cell shape.Our research focuseson the interphase corticalmicrotubulecytoskeleton becauseits organizational stateaffects cell shape andelongation by influencingthe patterning ofcell wall material. Ourgoal is to elucidate themolecular mechanismsthat govern local corticalmicrotubule alignmentas well as globalarray organization. The Dixit lab is alsointerested in the function of kinesin motorsin plants and in the role of motor proteindiversity in plant evolution. To address theseproblems, we are using a combination oflive-cell and single molecule imaging techniques,computational approaches, and thepowerful genetic tools of the model plantArabidopsis.Current research areas in the lab are:1. Identification and functional characterizationof the microtubule plus-endcomplex of plants.2. Developing a computer model to quantitativelystudy cortical microtubule organization.3. Structure-function studies of a group ofArabidopsis kinesins at the single moleculelevel.RECENT PUBLICATIONSDixit, R., Chang, E. and Cyr, R.J. (2006).Establishment of polarity during organizationof the acentrosomal plant cortical microtubulearray. Molecular Biology of the Cell,17: 1298-1305.Dixit, R., Cyr, R.J. and Gilroy, S. (2006).Using intrinsically fluorescent proteins forplant cell imaging. Plant Journal, 45: 599-615.Dixit, R. and Cyr, R.J. (2004). Encountersbetween dynamic cortical microtubules promoteordering of the cortical array throughangle-dependent modifications of microtubulebehavior. Plant Cell, 16: 3274-3284.Dixit, R. and Cyr, R.J. (2004). The corticalmicrotubule array: from dynamics to organization.Plant Cell, 16: 2546-2552.Dixit, R. and Cyr, R.J. (2003). Cell damage andreactive oxygen species production inducedby fluorescence microscopy: Effect on mitosisand guidelines for non-invasive fluorescencemicroscopy. Plant Journal, 36: 280-290.Dixit, R. and Cyr, R.J. (2002). Spatio-temporalrelationship between nuclear-envelopebreakdown and preprophase band disappearancein cultured tobacco cells. Protoplasma,219: 116-121.(A) Confocal image of cortical microtubules inArabidopsis epidermal cells. (B) Maximum projectionof pseudo-colored time-lapse images of EB1-GFPfrom a living tobacco BY2 cell. This image illustratesthe growth vector of individual cortical microtubulesgoing sequentially from blue to green to yellow to red.(C) Single GFP-labeled kinesin molecules (3 moleculescan be seen in this sequence) observed movingalong a rhodamine-labeled microtubule using totalinternal reflection fluorescence microscopy. The dottedline indicates the movement of one of the kinesinmolecules over 3 seconds.

Ian W. DuncanProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONWe are interested in how genes control thedevelopment of specific body segments inDrosophila. A major focus is the antennalsegment, since this develops independentlyof the Hox genes, which control the identitiesof most other body segments. The antennacontains both olfactory and auditory organs,and serves as both the nose and ear of thefly. We have shown that the distal (olfactory)part of the antenna is specified by the spineless(ss) gene, which encodes a transcriptionfactor that is homologous to the vertebratearyl hydrocarbon receptor. Much of our currentwork is devoted to understandinghow ss controls antennal development.We have been very successfulin identifying ss target genes bysearching the Drosophila genomesequence for clusters of binding sitesfor Ss and its coregulators. Thiswork promises to allow a globalunderstanding of the developmentalprogram for constructing an antenna;such an understanding has notbeen achieved for any other bodysegment or appendage type.Antennal specification by ss. (a) A normalantenna, showing two bristle-containingproximal segments (AI and AII),a bulbous third segment (AIII), whichserves as the major olfactory organ ofthe fly, and a distal plume-like arista(Ar). (b) Antenna from an ss mutantfly. The arista and part of AIII aretransformed to distal leg (tarsi). (c)Expression of an ss target gene.Staining of a lacZ reporter (blue) for thegene is seen specifically in AIII and thearista, where ss is expressed.We are also very interested in how patterningoccurs within body segments. For this work,we are focusing on the abdominal segmentsof the adult, which are much simpler thanthose in the thorax and head.SELECTED PUBLICATIONSEmmons, R.B., Duncan, D. and Duncan, I.(2007). Regulation of the Drosophila distalantennal determinant spineless.Developmental Biology, 302: 412-426.Wernet, M.F., Mazzoni, E.O., Celik, A.,Duncan, D.M., Duncan, I. and Desplan, C.(2006). Stochastic expression of theDrosophila dioxin receptor spineless createsthe retinal mosaic for color vision. Nature,440: 174-180.Kopp, A. and Duncan, I. (2002).Anteroposterior patterning in adult abdominalsegments of Drosophila. DevelopmentalBiology, 242: 15-30.Kopp, A., Duncan, I. and Carroll, S.B. (2000).Genetic control and evolution of sexuallydimorphic characters in Drosophila. Nature,408: 553-559.Duncan, D.M., Burgess, E.A., and Duncan, I.(1998). Control of distal antennal identityand tarsal development in Drosophila byspineless-aristapedia, a homolog of themammalian dioxin receptor. Genes andDevelopment, 12: 1290-1303.21

Sarah C.R. ElginProfessor of BiologyViktor Hamburger Professor in Arts & Scienceswww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONThe Elgin lab is interested in the role thatchromatin structure plays in gene regulation,considering both effects from packaging largedomains and local effects of the nucleosomearray. The lab works with Drosophilamelanogaster, the fruit fly, combining biochemical,genetic, and cytological approaches.It is investigating the mechanism of genesilencing associated with heterochromatinpackaging, looking both at the chromosomalproteins and histone modifications involved,and at the underlying DNA sequences thatare targets for heterochromatin formation.Possible RNAi-based mechanisms are beinginvestigated.SELECTED PUBLICATIONSGrewal, S.I.S. and Elgin, S.C.R. (2007).Transcription and RNA interference in theformation of heterochromatin. Nature, 447:399-406.Shaffer, C., Cenci, G., Thompson, B.,Stephens, G.E., Slawson, E., Adu-Wusu, K.,Gatti, M. and Elgin, S.C.R. (2006). The largeisoform of Drosophila melanogasterHeterochromatin Protein 2 plays a criticalrole in gene silencing and chromosomestructure. Genetics, 174: 1189-1204.Haynes, K.A., Caudy, A.A., Collins, L. andElgin S.C.R. (2006). Element 1360 and componentsof the RNAi system contribute toHP1-dependent silencing of a pericentricreporter. Current Biology, 16: 2222-2227.Sun, F.L., Haynes, K., Simpson, C.L., et al.(2004). cis-Acting determinants of heterochromatinformation on Drosophilamelanogaster chromosome four. MolecularCell Biology, 24: 8210-8220.Sun, F.L., Cuaycong, M.H., Elgin, S.C.R.(2001). Long-range nucleosome ordering isassociated with gene silencing in Drosophilapericentric heterochromatin. Molecular CellBiology 21: 2867-2879.Our P-element construct (based on vector A412 fromV. Pirrotta) contains a visible marker for variegation,hsp70-white, and a marked copy of hsp26 for subsequentstudies of chromatin structure. Several fly lineshave been recovered showing a PEV phenotype (A);all have P element inserts in the pericentric heterochromatin(as shown for this case by in situ hybridizationto the polytene chromosomes with the entire P element;see B), telomeres, or the fourth chromosome. (Note thatthe P element also hybridizes to the sites of the endogenouswhite, hsp70, and hsp26 genes.) See Wallrathand Elgin (1995) Genes & Develop 9: 1263-1277.22

Ursula W. GoodenoughProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONWe study the molecular basis and evolutionof sexual lifecycle transitions in the flagellatedgreen alga, Chlamydomonas reinhardtii.Current projects include: 1)Analysis of sex-determination, mediated bythe dominant Mid protein encoded in themating-type minus locus; 2) Analysis ofuniparental inheritance of plus chloroplastDNA, mediated by several genes encodedin the mating-type locus; 3) The haploiddiploidtransition, mediated by zygoticheterodimerization of homeoproteins contributedby plus and minus gametes; 4)Rapid evolution of sex-related genes, con-centrated in low-complexity regions of codingsequences. We also study hydroxyprolinerichglycoproteins in the cell wall and insexual flagellar agglutination.SELECTED PUBLICATIONSLee, J-H., Lin, H., Goodenough, U. (2007). AChalmydomonas KNOX protein, GSM1, initiateszygote differentiation with a BELL-relatedprotein, Gsp1. Manuscript submitted.Lee, J-H., Waffenschmidt, S., Small, L.,Goodenough, U. (2007). Between-speciesanalysis of short-repeat modules in cell-walland sex-related hydroxyproline-rich glycoproteinsof Chlamydomonas. PlantPhysiology, In Press.Lin, H., Goodenough, U. (2007).Gametogenesis in the Chlamydomonas reinhardtiiminus mating type is controlled bytwo genes, MID and MTD1. Genetics, InPress.Ferris, P.J., Waffenschmidt, S., Umen J.G.,Lin, H., Lee, J-H., Ischida, K., Kubo, T., Lau,J., Goodenough, U. (2005). Plus and minussexual agglutinins from Chlamydomonasreinhardtii. Plant Cell, 17: 597-615.Ferris, P.J., Armbrust, E.V., Goodenough, U.(2002). Genetic structure of the mating-typelocus of Chlamydomonas reinhardtii.Genetics, 160: 181-200.Two Chlamydomonasgametes, mating.23

Elizabeth S. HaswellAssistant Professor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTION:All organisms must sense and respond tomechanical forces generated outside the cell(shear force, gravity, touch) as well as insidethe cell (osmotic pressure, membrane deformation).The Haswell lab is interested in howphysical force is converted into a biochemicalsignal capable of altering the state of a cell.We are addressing this question in the modelplant Arabidopsis thaliana, where a numberof important mechanical signal transductionpathways have been characterized. We usean array of biochemical and molecular geneticapproaches, electrophysiology, and stateof-the-artlive imaging methodologies in ourexperiments.RECENT PUBLICATIONS:Haswell, E. S. (2007). MscS-like Proteinsin Plants. Current Topics in Membranes, 58:329-359.Haswell, E. S. and Meyerowitz, E.M. (2006).MscS-like Proteins Control Plastid Size andShape in Arabidopsis thaliana. CurrentBiology, 16: 1-11.Haswell, E. S. (2003). Gravity Perception:How Plants Stand up for Themselves. CurrentBiology, 13: R761-R763.Three main lines of inquiry in the lab are:1) functional characterization of a family ofmechanosensitive ion channels related to thebacterial channel MscS; 2) genetic approachesto identifying new components of gravityand touch signal transduction pathways;and 3) investigation into the role played bymechanosensory systems in organelle shapeand size determination.(A) Plastids from a plant expressing mutantmechanosensitive ion channels have an unusualspherical morphology (the normal football-shapedplastid found in a wild-type plant is shown inthe inset). Plastids are plant-specific organellesresponsible for photosynthesis, gravity response,and a number of important metabolic reactions.(B) Versions of these channels fused to GreenFluorescent Protein localize in clusters (shownin green) at the poles of the plastids. We wouldlike to know why mechanosensitive channels areclustered at the plastid poles, and how they contributeto plastid shape and size.24

Erik D. HerzogAssociate Professor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONBiological clocks that drive daily rhythms inbehavior and physiology have been found ina wide variety of organisms and cell types.The Herzog lab studies the cellular andmolecular basis of these circadian rhythms inmammals using techniques including multielectrodearrays, cellular imaging, pharmacologicaland genetic manipulations, andbehavioral assays. This approach is producinginsight into the roles of specific molecules,neurons, glia, and brain areas in coordinatingour daily schedules.For example, the suprachiasmatic nucleus(SCN) of the hypothalamus is a master circadianpacemaker. This brain region continuesto keep daily time when cultured. We arenow asking which cells generate this rhythmicity,how do they synchronize to oneanother, and how do they send their timinginformation to the rest of the brain and body?SELECTED PUBLICATIONSAton, S.J., Huettner, J.E., Straume, M.,Herzog, E.D. (2006). GABA and Gi/o differentiallycontrol circadian rhythms and synchronyin clock neurons. Proceedings of theNational Academy of Sciences USA, 103:19188-19193.Granados-Fuentes, D., Tseng, A., Herzog,E.D. (2006). A circadian clock in the olfactorybulb controls olfactory responsivity.Journal of Neuroscience, 26: 12219-12225.Herzog, E.D., Muglia, L.J. (2006). You arewhen you eat. Nature Neuroscience, 9: 300-302.Aton, S.J., Herzog, E.D. (2005). Cometogether, right...now: synchronization ofrhythms in a mammalian circadian clock.Neuron, 48: 531-534.Prolo, L.M., Takahashi, J.S., Herzog, E.D.(2005). Circadian rhythm generation andentrainment in astrocytes. Journal ofNeuroscience, 25: 404-408.Clocks in a dish. Representative firing patterns of three rhythmicSCN neurons recorded on neighboring electrodes (left plots). Pairwisecross-correlations of the discharge rate binned at 6-minintervals show a significant peak at 0 h lag on a 24-h timescaleindicating circadian synchrony (middle). Grey lines show the 95percent confidence interval derived by randomizing the neurons’firing patterns and cross-correlating 1000 times. The same neuronpairs failed to show spike-for-spike cross-correlations at any circadiantime (right). Spike-for-spike correlations are shown for10 min of data collected at 240 h (arrows on left).25

Tuan-hua David HoProfessor of Biologywww.biology.wustl.edu/facultyCover of Plant Physiology(November issue, 2002)highlighting the study ofHVA22 homolog in yeastfollowing yeast syntheticlethal mutant screening.RESEARCH DESCRIPTIONHormone and Stress-Regulated GeneExpression in PlantsThe Ho lab is interested in mechanismsregulating the developmental transitionfrom embryogenesis to seedgermination, a process governedby two phytohormones, gibberellins(GA) and abscisic acid (ABA). Sinceplant embryos are very resistant toenvironmental stresses such asdrought and cold, they also studythe role of stress-induced proteinsin mediating stress tolerance inplants.Hormonal Control of GeneExpression in the Aleurone Cellsof Cereal GrainsIn this project the Ho lab studies theeffect of GA and ABA on the expressionof genes encoding alpha-amylases,proteases, and nucleases ingerminating cereal grains. They followboth biochemical and geneticapproaches to investigate the functionof genes involved in siginalingprocess. In particular, the Ho labplans to investigate the interactionsamong three regulatory factors,GAMyb, SLN, and PKABA1.tein, HVA22, has apparent homologs inmany diverse eukaryotes but not in anyprokaryotes. The lab’s current results suggestthat HVA22 is likely involved in vesiculartrafficking and autophagy.SELECTED PUBLICATIONSCasaretto, J. A. and Ho, T-H. D. (2005).Transcriptional regulation by abscisic acid inbarley (Hordeum vulgare L.) seeds involvesautoregulation of the transcription factorHvABI5. Plant Molecular Biology, 57: 21-34.Casaretto, J. and Ho, T-H. D. (2003). TheTranscription Factors HvABI5 and HvVP1Are Required for the Abscisic Acid Inductionof Gene Expression in Barley Aleurone Cells.Plant Cell, 15: 271-284.Brands, A. and Ho, T-H. D. (2002). Functionof a Plant Stress-Induced Gene, HVA22.Synthetic Enhancement Screen with Its YeastHomolog Reveals Its Role in VesicularTraffic1. Plant Physiology, 130: 1121-1131.Zentella, R., Yamauchi, D. and Ho, T-H. D.(2002). Molecular Dissection of theGibberellin/Abscisic Acid Signaling Pathwaysby Transiently Expressed RNA Interference inBarley Aleurone Cells. Plant Cell, 14: 2289-2301.Cover of Plant Cell(January issue, 2003)highlighting the use ofcereal aleurone system ininvestigating hormonalregulation.26Function of Stress-Induced GenesThe Ho lab is particularly interestedin the function of drought, salinity,and cold stress induced proteins incereals and Arabidopsis. One ofthese proteins, HVA1, contains longstretches of amphipathic alpha-helicalstructure, and its over-expressionin transgenic plants leads toelevated levels of stress tolerance.Another stress/ABA-induced pro-Gomez-Cadenas, A., Zentella, R., Walker-Simmons, M. K. and Ho, T-H. D. (2001).Gibberellin/Abscisic Acid Antagonism inBarley Aleurone Cells. Site of action of theprotein kinase PKABA1 in relation to gibberellinsignaling molecules. Plant Cell, 13:667-679.

Tiffany M. KnightAssistant Professor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONThe Knight lab studies the population ecologyof rare and invasive plant species andaddresses questions related to the causes andconsequences of their abundances and distributions.Why are some species rare, whiletheir closely related congeners are widespread?How does dispersal ability and densitydependence determine the populationabundance and spread of invasive plants?In particular, Knight focuses on the role ofinterspecific interactions between plants andherbivores, seed predators, pollinators, andcompetitors on the long-term rates of plantpopulation growth. She considers howinbreeding and inbreeding depression mayaffect interspecific interactions. Her studysites include the Tyson Research Centerand the Shaw Nature Reserve (both nearSt. Louis, Missouri) and Point ReyesNational Seashore in Northern California.RECENT PUBLICATIONSKnight, T.M., Steets, J.A. and Ashman, T-L.(2006). A quantitative synthesis of pollensupplementation experiments highlights thecontribution of resource reallocation to estimatesof pollen limitation. American Journalof Botany, 92: 270-276.Knight, T.M., McCoy, M.W., Chase, J.M.,McCoy, K.A. and Holt, R.D. (2005). Trophiccascades across ecosystems. Nature, 437:880-883.Knight, T.M., Steets, J.A., Vamosi, J.C. et al.(2005). Pollen limitation of plant reproduction:Ecological and Evolutionary Causes andConsequences. Annual Review of Ecology,Evolution and Systematics, 36: 467-497.Knight, T.M. (2004). The effects of herbivoryand pollen limitation on a declining populationof Trillium grandiflorum. EcologicalApplications, 14: 915-928.Photos show the widespread species, Lupinuschamissonis (top) and its co-occurring congener,Lupinus tidestromii (bottom), which is rare andfederally endangered. Differences between thesespecies in their levels of seed predation by rodentscause L. chamissonis to have significantly higherrates of population growth than L. tidestromii.27

Robert G. KranzProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONThe Kranz group studies two differentprocesses using various model organismsthat are best suited for each process:i. Cytochrome c biogenesis pathways (seethe figure). Using biochemical, genetic, andimmunological approaches, the group studiesa photosynthetic bacterium, Rhodobactercapsulatus, for System I and Bordetella pertussisas a model for System II. RecombinantE. coli with all three systems, including thehuman system III, are also investigated.Since c-type cytochromes are requiredfor growth in most organisms, as well asapoptosis in eukaryotes and as bacterialnanowires, understanding and controllinghow they are synthesized is essential.ii. The control of gene expression in responseto nitrogen in the environment, using R.capsularus and Arabidopsis.SELECTED PUBLICATIONSRichard-Fogal, C.L., Frawley, E.R., Feissner,R.E. and Kranz, R.G. (2007). Heme concentrationdependence and metalloporphyrininhibition of the system I and II cytochrome cassembly pathways. Journal of Bacteriology,189(2): 455-463.Engineer, C. B. and Kranz, R.G. (2006).Reciprocal leaf and root expression ofAtAmt1.1 and root architectural changesin response to nitrogen starvation. PlantPhysiology, 143(1): 236-250.Feissner R.E., Richard-Fogal, C.L., Frawley,E.R., Kranz, R.G. (2006). ABC transportermediatedrelease of a heme chaperoneallows cytochrome c biogenesis. MolecularMicrobiology, 61(1): 219-231Feissner R.E., Richard-Fogal, C.L., Frawley,E.R., Loughman, J.A., Earley, K.W., Kranz,R.G. (2006). Recombinant cytochromes cbiogenesis systems I and II and analysis ofhaem delivery pathways in Escherichia coli.Molecular Microbiology, 60(3): 563-577.Feissner, R.E., Beckett, C.S., Loughman, J.A.,Kranz, R.G. (2005). Mutations in cytochromeassembly and periplasmic redox pathways inBordetella pertussis. Journal of Bacterioogy,187(12): 3941-3949.Three unique systems have evolved to assemble thec-type cytochromosomes. The assembly proteins foreach system are shown. (c-type cytochromes haveheme that is covalently linked to two cysteines.)28

Barbara N. KunkelAssociate Professor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONBarbara Kunkel’s research group studies theregulatory mechanisms governing interactionsbetween bacterial plant pathogens andtheir hosts. Specifically, they would like tounderstand the molecular basis of pathogenesisand disease development and are focusingon the identification and characterizationof pathogen virulence factors and their targetsin plant cells. The group studies the bacterialplant pathogen Pseudomonas syringaeand two of its hosts, Arabidopsis thalianaand tomato, systems in which both pathogenand host are amenable to genetic and molecularanalyses. Kunkel’s research currentlyfocuses on two central questions: 1) Whatis the molecular basis of pathogenesis in P.syringae, and 2) What signaling pathwaysand physiological processes within the hostare manipulated by the pathogen duringinfection?SELECTED PUBLICATIONSLaurie-Berry, N., Joardar, V., Street, I.H.and Kunkel, B.N. (2006). The Arabidopsisthaliana JASMONATE INSENSITIVE 1 geneis required for suppression of salicylic aciddependentdefenses during infection byPseudomonas syringae. Molecular Plant-Microbe Interactions, 19: 789-800.Preiter, K., Brooks, D.M., Penaloza-Vazquez,A., Sreedharan, A., Bender, C.L. and Kunkel,B.N. (2005). Novel virulence gene ofPseudomonas syringae pathovar tomatostrain DC3000. Journal of Bacteriology, 187:7805-7814.Brooks, D.M., Bender, C.L. and Kunkel, B.N.(2005). The Pseudomonas syringae phytotoxincoronatine promotes virulence by overcomingsalicylic acid-dependent defenses inArabidopsis thaliana. Molecular PlantPathology, 6: 629-639.Lim, M.T.S. and Kunkel, B.N. (2005). ThePseudomonas syringae gene avrRpt2 contributesto virulence on tomato. MolecularPlant-Microbe Interactions, 18: 626-633.The phytotoxin coronatine is a key virulence factorfor the bacterial plant pathogen Pseudomonassyringae. Arabidopsis plants infected with wildtype(WT) P. syringae develop disease, whereasplants infected with a coronatine biosynthetic(COR - ) mutant do not. Coronatine, a molecularmimic of the plant hormone jasmonate, is criticalat multiple stages during P. syringae pathogenesis,including: 1) facilitating entry into leaf tissueby stimulating opening of stomata, 2) inhibitinghost defenses that restrict pathogen growth, and3) promoting disease symptom development. Fordetails, see Brooks et al, 2005.Lim, M.T.S. and Kunkel, B.N. (2004). ThePseudomonas syringae type III effectorAvrRpt2 promotes virulence independentlyof RIN4, a predicted virulence target inArabidopsis thaliana. Plant Journal, 40:790-798.Chen, Z., Kloek, A.P., Cuzick, A.,Tang, D.,Moeder, W., Klessig, D., McDowell, J.,Innes, R. and Kunkel, B.N. (2004). ThePseudomonas syringae AvrRpt2 proteinfunctions downstream or independentlyof salicylic acid to promote virulence onArabidopsis thaliana. Plant Journal, 37:494-504.29

Allan LarsonProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONAllan Larson’s research combines naturalhistory and molecular genetics to examinepatterns and processes of evolution at multiplelevels of biological complexity. Molecularphylogenetics provides a framework for integratingknowledge of historical biogeography,speciation, genome evolution, anddevelopmental/morphological evolution atboth macroevolutionary and microevolutionarytimescales. Salamanders and lizards providethe main study systems, with particularattention in the latter group to comparativephylogeography of continental and islandforms. Graduate-level teaching includescourses in macroevolution,molecular evolution,and history ofevolutionary theory.SELECTED PUBLICATIONSKozak, K.H., Blaine, R.A. and Larson, A.(2006). Gene lineages and eastern NorthAmerican palaeodrainage basins: phylogeographyand speciation in salamanders ofthe Eurycea bislineata species complex.Molecular Ecology, 15: 191-207.Kozak, K.H., Weisrock, D.W. and Larson, A.(2006). Rapid lineage accumulation in anon-adaptive radiation: phylogeographicanalysis of diversification rates in easternNorth American woodland salamanders(Plethodontidae: Plethodon). Proceedings ofthe Royal Society of London, Series B, 273:539-546.Nicholson, K.E., Mijares-Urrutia, A. andLarson, A. (2006). Molecular phylogeneticsof the Anolis onca series: a case history inretrograde evolution revisited. Journal ofExperimental Zoology (Molecular andDevelopmental Evolution), 306B: 450-459.Weisrock, D.W. and Larson, A. (2006).Testing hypotheses of population-level lineagesin the Plethodon jordani species complexwith allozyme and mitochondrial DNAsequence data. Biological Journal of theLinnean Society, 89: 25-51.An Asian salamander,Tylototriton, shanjing.Weisrock, D.W., Papenfuss, T.J., Macey, J.R.,Litvinchuk, S.N., Polymeni, R., Ugurtas, I.H.,Zhao, E., Jowkar, H. and Larson, A. (2006).A molecular assessment of phylogenetic relationshipsand lineage accumulation rateswithin the family Salamandridae (Amphibia,Caudata). Molecular Phylogenetics andEvolution, 41: 368-383.30

Petra Anne LevinAssistant Professor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONHow cells determine when and where todivide remains one of the great mysteries ofmodern biology. Spatially, division is tightlyregulated to ensure the accurate positioningof a division septum within individual cells.Temporally, division is coordinated with DNAreplication and chromosome segregation toguarantee that each daughter cell receives acomplete genome.Taking a broad-based approach that incorporatesgenetics, cell biology, and biochemistrythe Levin lab’s research seeks to characterizethe regulatory networks governing cell divisionin the Gram-positive bacterium Bacillussubtilis. By unraveling this process in a relativelysimple, tractable model system, we willfurther our understanding not only of bacterialcytokinesis but also of the fundamentalmechanisms responsible for the temporaland spatial regulation of cell division in allorganisms.SELECTED PUBLICATIONSHaeusser, D.P., Garza, A.C., Buscher, A.Z.and Levin, P.A. (2007). The divisioninhibitor EzrA contains a seven-residuepatch required for maintaining the dynamicnature of the medial FtsZ ring. Journal ofBacteriology, 189: 9001-9010.Weart, R. B., Lee, A., Chien, A-C., Haeusser,D.P., Hill, N.S. and Levin, P.A. (2007). Ametabolic sensor governing cell size in bacteria.Cell, 130: 335-347.Weart, R. B., Nakano, S., Lane, B.E., Zuber,P. and Levin, P.A. (2005). The ClpX chaperonemodulates assembly of the tubulin-likeprotein FtsZ. Molecular Microbiology, 57:238-249.Haeusser, D.P., Schwartz, R.L., Smith, A.M.,Oates, M.E. and Levin, P.A. (2004). EzrAprevents aberrant cell division by modulatingassembly of the cytoskeletal protein FtsZ.Molecular Microbiology, 52: 801-814.False colored images of the B. subtilis cellsstained for the cell division protein FtsZand DNA. FtsZ forms a distinctive band atthe middle of each cell in these images.31

Kathryn G. MillerProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONSELECTED PUBLICATIONS1.2.Multicellular organisms have many differentcell types that are specialized to have rolesimportant to the normal functioning of theorganism. Generating and maintaining thespecialized morphologies and functionalproperties of these cells usually depends onactin cytoskeletal organization. To understandthe roles of actin cytoskeletal structuresin some of these specialized cell types,the Miller lab uses genetic and moleculargenetic techniques to alter the function ofproteins of the actin cytoskeleton. Thesestudies use the model system, Drosophila,because this type of functional manipulationis relatively straightforward. Examination ofresulting disruptions of cell morphology andfunction using cell biological, biochemical,and imaging methods both in vitro and invivo (using GFP and other probes), permit usto understand contributions of different actinregulating and actin-associated proteins tocell function. In addition, these studies illustratethe general functionsof the many different typesof actin structures presentin cells. Because actin andits associated proteins arehighly conserved acrossall eukaryotic species, theinformation we obtainabout mechanism of actincytoskeletal organizationand function is widelyapplicable.Frank, D.J., Hopmann, R., Lenartowska, M.,Miller, K.G. (2006). “Capping Protein and theArp2/3 Complex Regulate Non-Bundle ActinFilament Assembly to Indirectly ControlActin Bundle Positioning during Drosophilamelanogaster Bristle Development.” MolecularBiolology of the Cell, 17(9): 3930-3939.Frank, D.J., Martin, S.R., Gruender, B.N., Lee,Y.S., Simonette, R.A., Bayley, P.M., Miller,K.G., Beckingham, K.M. (2006). “Androcamis a tissue-specific light chain for myosinVI in the Drosophila testis.” Journal ofBiological Chemistry, 281(34): 24728-24736.Noguchi, T., Lenartowska, M. and Miller, K.G.(2006). Myosin VI stabilizes a branchedactin meshwork during spermatid individualizationin Drosophila. Molecular Biology ofthe Cell, 17: 2559.Noguchi, T. and Miller, K.G. (2003). A rolefor actin dynamics in individualizationduring spermatogenesis in Drosophila.Development, 130: 1805-1816.Figure 1: Actin Cones and Myosin VIFluorescence image of spermatids during the individualizationstage of Drosophila spermatogenesis.Actin structures called actin cones (red) areimportant for membrane reorganization duringsperm development in Drosophila. Myosin VI, anactin-based motor protein (green), helps stabilizethis structure. The cones move from the base ofthe spermatid nuclei (blue) for over 2 microns tothe end of the axoneme. In this image, the coneshave moved only slightly away from the nuclei.32Figure 2: Developing Pupal BristlesA fluorescence image of a developing Drosophilabristle cell. Actin bundles (red) help shape thebristle and small patches of dynamic actin calledactin snarls (in this case labeled to reveal anactin assembly regulator; green) lie between thebundles. The snarls help to position the bundlesas the bristle grows.

Kenneth M. OlsenAssistant Professor of Biologywww.biology.wustl.edu/faculty1.RESEARCH DESCRIPTION:The Olsen lab’s research focuses on thegenetic basis of evolution in plants: how isthe variation that we find within a speciesshaped by natural selection, population history,and other evolutionary forces? One waythat we look at this question is by using cropdomestication as a genetic model for rapidevolutionary change. Currently, we arestudying the evolutionary genomics of weedyrice (red rice), which is a major pest in U.S.rice fields. In addition to crop domestication,our lab also studies the evolutionary geneticsof wild plant species. Topics of interestinclude the genetic basis of adaptive variation,the forces affecting genome-wide patternsof linkage disequilibrium, and phylogeography.A current research focus is on themolecular evolution of an adaptive polymorphismfor cyanogenesis (cyanide production)in white clover.RECENT PUBLICATIONS:Olsen, K.M., Sutherland, B.L. and Small, L.L.(2007). Molecular evolution of the Li/Lichemical defence polymorphism in whiteclover (Trifolium repens L.). MolecularEcology, 16: 4180-4193.Olsen, K.M., Caicedo, A.L. and Jia, Y. (2007).Evolutionary genomics of weedy red rice inthe USA. Journal of Integrative PlantBiology, 49(6): 811-816.Narasimhamoorthy, B., Bouton, J.H., Olsen,K.M. and Sledge, M.K. (2007). Quantitativetrait loci and candidate gene mapping ofaluminum tolerance in diploid alfalfa.Theoretical and Applied Genetics, 114:901–913.Olsen, K.M., Caicedo, A.L., Palato, N.,McClung, A., McCouch, S. and Purugganan,M.D. (2006). Selection under domestication:evidence for a sweep in the rice Waxygenomic region. Genetics, 173: 975-983.Kuo, H-F., Olsen, K.M. and Richards, E.J.(2006). Natural variation in a subtelomericregion of Arabidopsis: implications for thegenomic dynamics of a chromosome end.Genetics, 173: 401-417.2.Figure 1: White clover (Trifolium repens) isnaturally polymorphic for cyanogenesis (cyaniderelease with tissue damage). Our lab is studyingthe molecular evolution of this adaptive polymorphism.Figure 2: Seeds from different strains of red rice(Oryza sativa). This weedy relative of rice infestsagricultural fields and can reduce crop harvestsby 80 percent. We are studying the evolutionarygenomics of this weed.33

John L. OrrockAssistant Professor of Biologywww.biology.wustl.edu/facultyAn experimental exclosurein the grasslandsof California to measureconsumer impacton plant communities(photo by EllenDamschen); Inset: adeer mouse leaves aforaging tray on SanMiguel Island (photoby John Orrock).34RESEARCH DESCRIPTIONWork in the Orrock lab centers around threethemes: behavior, ecological interactions,and how space mediates ecological and evolutionarydynamics. Our work is particularlyfocused on questions where these threethemes converge. For example, are invasiveplants successful because they provide apredator-free refuge from which nativeconsumers eat native plants?Although conservation corridorsbenefit plants by increasing seeddispersal, are these benefits offsetby corridor-mediated changes inthe foraging behavior of seed-eatingconsumers? The work we dois both basic and applied: Wecombine behavioral and spatialecology to provide insight into the forcesshaping communities, and thus also providea way to understand the ecological implicationsof rapid changes in landscape composition(e.g. by humans or invasive organisms).Research sites include the grasslands ofCalifornia, oldfields in South Carolina,Missouri oak forests, and the ChannelIslands off the California coast.Current Projects1. Apparent competition and invasiveplants: understanding how changes inrodent behavior and abundance causedby exotic plants might facilitate theirinvasion.2. Trophic cascades and the evolution ofanti-predator behavior in insular systems.3. The comparative role of consumptive andnon-consumptive effects: how predatorsalter prey dynamics without killing prey.4. Evaluating the role of patch shape andconnectivity in mediating the effect ofconsumers on plant communities.RECENT PUBLICATIONSDamschen, E.I., Haddad, N.M., Orrock, J.L.,Levey, D.J. and Tewksbury, J.J. (2006).Corridors increase plant species richness atlarge scales. Science, 313: 1284-1286.Orrock, John L., Levey, D.J., Danielson, B.J.and Damschen, E.I. (2006). Seed predation,not seed dispersal, explains the landscapelevelabundance of an early-successionalplant. Journal of Ecology, 94: 838-845.Orrock, John L. and Fletcher, Jr., R.L. (2005)Changes in community size affect the outcomeof competition. American Naturalist,166: 107-111.Orrock, John L. and Damschen, E.I. (2005).Corridors cause differential seed predation.Ecological Applications, 15: 793-798.Orrock, John L., Danielson, B.J. andBrinkerhoff, J. (2004). Rodent foraging isaffected by indirect, but not by direct, cuesof predation risk. Behavioral Ecology, 15:433-437.Orrock, John L. and Danielson, B.J. (2004).Rodents balancing a variety of risks: invasivefire ants and indirect and direct indicatorsof predation risk. Oecologia, 140: 662-667.Orrock, John L., Danielson, B.J., Burns, M.J.and Levey, D.J. (2003) Spatial ecology ofpredator-prey interactions: corridors andpatch shape influence seed predation.Ecology, 84: 2589-2599.

Philip A. OsdobyProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONThe cells and matrix comprising skeletal tissueform a specialized architecture uniquelyadapted for weight bearing, motion, andmarrow development. Bone also serves as amineral reservoir for the organism and is, bynecessity, highly responsive to complicatedendocrine signals responsible for maintainingand modifying mineral homeostasis. In addition,the complex bone matrix may serve as agrowth factor repository and thereby operateto influence haematopoietic events and contributeto the regenerative properties of bone.Bone modeling during development andremodeling throughout life are dependentupon factors that regulate the number andactivity of both bone-forming osteoblasts andbone-resorbing multinucleated osteoclasts.Deciphering the biochemical and molecularmechanisms that govern bone cell precursorrecruitment, differentiation, and activity is,therefore, important for understanding bothnormal and pathological processes in bone.The osteoclast is believed to degrade bonematrix by releasing hydrolytic enzymes,superoxide radicals, and protons into whatcan best be described as an extracellularphagolysosome. There are unanswered questionsabout the developmental relationshipbetween osteoclasts (OC), OC precursors. Theresearch in our group focuses on: identifyingand characterizing unique osteoclast plasmamembrane proteins to better understandinghow the specialized bone-resorbing ability ofthis cell is accomplished and regulated; identifyingfactors derived from the bone environmentthat influence osteoclast developmentand determining how these signalsmay be modified during development andaging; examining the functional and regulatoryrole of Nitric oxide (NO) in bone remodelingincluding studies designed to use NOtherapy to alleviate the bone fragility associatedwith osteogenesis Imperfecta. Projectalso centers on examining the role ofchemokines, e.g., SDF-1 in bone remodelingand fracture repair.SELECTED PUBLICATIONSZheng, H., Yu, H., Collin-Osdoby, P., Osdoby,P. (2006). RANKL Stimulates iNOS and NOproduction in Developing Osteoclasts viaNF-kB: An Autocrine Negative FeedbackMechanism to Regulate Osteoclastogenesisand Bone Resorption. Journal of BiologicalChemistry, 281(23): 15809-15820.Kindle, L., Rothe, L., Kriss, M., Osdoby, P.,Collin-Osdoby, P. (2006). Human microvascularendothelial cell activation by IL-1and TNF-a stimulates the adhesion andtransendothelial migration of circulatinghuman CD14+ monocytes that develop withRANKL into functional osteoclasts. Journal ofBone and Mineral Resources 21: 193-206.Yu, X., Huang, Y., Collin-Osdoby, P., Osdoby,P. (2003). Stromal cell-derived factor-1(SDF-1) recruit osteoclast precursors by inducingchemotaxis, matrix metalloproteinase-9(MMP-9) activity, and collagen transmigration.Journal of Bone and Mineral Research,18: 1404-1418. 35

Himadri B. PakrasiDirector, International Center for Advanced Renewable Energy and Sustainability;George William and Irene Koechig Freiberg Professor of Biology; Professor ofEnergy, School of Engineeringwww.biology.wustl.edu/faculty36RESEARCH DESCRIPTIONThe current research in the lab focuses onapplication of systems biology approachesin the general area of bioenergy. The centralapproach in most of the biological sciencesduring past the 50 years has been based onreductionism and has resulted in a massiveincrease in our knowledge of individual cellularcomponents. However, we have alsorealized that while we know how the individual“nuts and bolts” function in isolation,our understanding of how these componentsinteract with each other to define the cellular,organismal, and population level behaviorsof living beings has remained far lesssophisticated. A “Systems Biology” approachshould provide enabling technologies toexamine complex biological processes, whichshould in turn result in an integrated andpredictive understanding of how an organismbehaves and responds to environmentalchanges. We are using systems approachesto determine the underlying network thatgoverns photosynthetic processes incyanobacteria, vascular plants (Arabidopsis),and mosses. In photosynthetic organisms,the cellular components are always in flux,CO 25C SugarADPCARBOXYLATIONREGENERATIONATPC 33C SugarCycleChloroplast StromaTriose-PREDUCTIONATPStarchNADP + + H + NADPHSynthesisFdH + FNRH +PQP PQhPSIPQHPSII2Cyt bfPCPCH +2 H 2O O 2+ 4H +ThylakoidMembraneSucroseSynthesisPQH 2CytoplasmThylakoid LumenPhotosynthesis in Chloroplasts.ATPSynthaseH +Chloroplast EnvelopeMembranesADP + P iH +and molecular machines assemble,function, and disassemble as a functionof time and environmental alterations suchas light intensity and nutrient availability.It is imperative to utilize a systems biologyapproach and integrate temporal informationfrom transcriptome, proteome, and metabolomelevel studies into a predictive,dynamic model to understand the functioningof a photosynthetic organism. We areleading two large-scale, multi-institutional,systems biology projects to pursue this goal.SELECTED PUBLICATIONSRoose, J.L., Kashino, Y. and Pakrasi, H.B.(2007). The PsbQ Protein DefinesCyanobacterial Photosystem II Complexeswith Highest Activity and Stability.Proceedings of the National Academy ofSciences USA, 104: 2548-2553.Aurora, R., Hihara, Y. and Pakrasi, H.B.(2007). A Network of Genes Regulated byLight in Cyanobacteria. OMICS, 11(2): 166-185.Koropatkin, N.M., Pakrasi, H.B. and Smith,T.J. (2006). Atomic structure of a nitratebindingprotein crucial for photosyntheticproductivity. Proceedings of the NationalAcademy of Sciences USA, 103: 9820-9825.Keren, N., Ohkawa, H., Welsh, E.A., Liberton,M. and Pakrasi, H.B. (2005). Psb29, aConserved 22-kD Protein, Functions in theBiogenesis of Photosystem II Complexes inSynechocystis and Arabidopsis. Plant Cell, 17:2768-2781.Thornton, L.E., Ohkawa, H., Roose, J.L.,Kashino, Y., Keren, N. and Pakrasi, H.B.(2004). Homologs of Plant PsbP and PsbQProteins are Necessary for Regulation ofPhotosystem II Activity in the Cyanobacterium,Synechocystis 6803. Plant Cell, 16: 2164-2175.

Craig S. PikaardProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONIn most offspring there are genes that areexpressed from the chromosomes inheritedfrom only one parent. Often a maternal orpaternal imprint dictates which allele will beactive. However, this is not the case for theuniparental expression of ribosomal RNA(rRNA) genes in genetic hybrids. This epigeneticphenomenon, known as nucleolar dominance,occurs both in plants and animals.The Pikaard lab has shown that rRNA genesilencing involves concerted changes in bothDNA methylation and histone modificationand has proposed a model whereby DNA andhistone modifications are each upstream ofone another in a self-reinforcing, circularpathway. Members of the lab are identifyingthe chromatin modifying activities involvedin this repression cycle. Two histone deacetylases,one DNA methyltransferase, and twomethylcytosine binding proteins have beenidentified in the screens thus far. They areusing genetic, cytogenetic, and biochemicalapproaches in order to understand how theactivities of these chromatin modifying activitiesare intertwined to comprise an epigeneticon-off switch. A second emphasis in thePikaard lab is the characterization of nuclearRNA polymerase IV (pol IV), whose atypicalcatalytic subunits were first identified byPikaard during the annotation of theArabidopsis genome sequence in 2000. Thereare two forms of Pol IV which play nonredundantroles in the production of smallinterfering RNAs (siRNAs) and the subsequentsiRNA-directed DNA methylation andsilencing of repeated genomic sequences,including transposable elements. Acting in apathway that includes DICER-LIKE 3 (DCL3),ARGONAUTE 4 (AGO4) and RNA-DEPEN-DENT RNA POLYMERASE 2 (RDR2), pol IVfunction is required to maintain the normalorganization of the chromatin within thenucleus. By combining genetic and cytologi-cal analyses, the Pikaard lab has determinedthe order of events in the Pol IV-dependentnuclear siRNA pathway and defined a siRNAprocessing center located in the nucleolus.Current priorities include purifying pol IVaand Pol IVb to homogeneity, determiningtheir complete subunit structures, determiningtheir template requirements (e.g. DNA vs.RNA), and identifying their transcripts orenzymatic products.SELECTED PUBLICATIONSEarley, Keith, Lawrence, Richard J., Pontes,Olga, Reuther, Rachel, Enciso, Angel J., Silva,Manuela, Neves, Nuno, Gross, Michael,Viegas, Wanda and Pikaard, Craig S.,(2006). Erasure of histone acetylation byArabidopsis HDA6 mediates large-scale genesilencing in nucleolar dominance. Genes &Development, 20: 1283-1293.Pontes, Olga, Li, Carey Fei, Nunes, PedroCosta, Haag, Jeremy, Ream, Thomas, Vitins,Alexa, Jacobsen, Steven E. and Pikaard, CraigS. (2006). The Arabidopsis chromatin-modifyingnuclear siRNA pathway involves anucleolar RNA processing center. Cell, 126:79-92.Onodera, Yasuyuki, Haag, Jeremy, Ream,Thomas, Nunes, Pedro Costa, Pontes, Olgaand Pikaard, Craig S. (2005). Plant NuclearRNA polymerase IV mediates siRNA andDNA methylation-dependent heterochromatinformation. Cell, 120: 613-622.Lawrence, R.J., Earley, K., Pontes, O., Silva,M., Chen, Z.J., Neves, N., Viegas, W.,Pikaard, C.S. (2004). A concerted DNAmethylation/histone methylation switch regulatesrRNA gene dosage control and nucleolardominance. Molecular Cell, 13: 599-609.37

Ralph S. QuatranoSpencer T. Olin Professor and ChairmanDepartment of Biologywww.biology.wustl.edu/faculty38RESEARCH DESCRIPTIONRalph S. Quatrano is interested in the mechanismsunderlying how cells become polar andhow tissue-specific factors and hormones regulategene expression in plants. The moss(Physcomitrella patens) is being used(http://www.biology.wustl.edu/moss) to studycellular polarity and homologous recombination,while Arabidopsis and P. patens are themodels for analyzing tissue-specific geneexpression via the phytohormone abscisicacid (ABA), specifically with respect todrought and desiccation tolerance. A comparativegenomic approach is also under waywith homologous genes that are part of anABA response pathway that has been conservedbetween an early land plant (i.e.moss) and seed plants (~450 million years).When tip growing moss filaments are givenan orienting gradient (e.g. light, gravity),what are the downstream targets for thesignaling path to direct polar growth? Ourhypothesis is that since the actin cytoskeletonhas been an essential and central link in ourunderstanding of polar processes in plants,the protein complexes that regulate the actinnetwork are the targets for signals that governpolar growth. These targets can also helpto identify interacting proteins that may localizeand stabilize these complexes to the polarsite. Hence, our directed approach is to focuson members of the Arp2/3 and theWave/SCAR protein complexes that regulateactin filament formation in otherorganisms. We are also employing a forwardgenetics approach using insertionalmutagenesis and activation tagging toTwenty-two-day-old Physcomitrella culture(see Cove, et al. (2006) Ann. Rev. PlantBiol. 57: 497-520.identify genes that affect polarized growth.These projects will be greatly aided by thecomplete sequence of the moss genome(http://www.mossgenome.org/).SELECTED PUBLICATIONSKhandelwal, A., Chandu, D., Roe, C.M.,Kopan, R. and Quatrano, R.S. (2007).Moonlighting activity of presenilin in plantsis independent of γ-secretase and evolutionarilyconserved. Proceedings of the NationalAcademy of Sciences USA, 104: 13337-13342.Quatrano, R.S., McDaniel, S.F., Khandelwal,A., Perroud, P-F. and Cove, D.J. (2007).Physcomitrella patens: mosses enter thegenomic age. Current Opinion in PlantBiology, 10: 182-189.Marella, H., Sakata, Y. and Quatrano, R.S.(2006). Characterization and functionalanalysis of ABSCISIC ACID INSENSITIVE3-like genes from Physcomitrella patens. ThePlant Journal, 46: 1032-1044.Perroud, P-F. and Quatrano, R.S. (2006). Therole of ARPC4 in tip growth and alignment ofthe polar axis in filaments of Physcomitrellapatens. Cell Motility and the Cytoskeleton, 63:162-171.Harries, P., Pan, A., and Quatrano, R.S.(2005). Actin-related protein2/3 complexcomponent ARPC1 is required for proper cellmorphogenesis and polarized cell growth inPhyscomitrella patens. The Plant Cell, 17:2327-2339.Lee, K.J.D., Sakata, Y., Mau, S-L., Pettolino,F., Bacic, A., Quatrano, R.S., Knight, C.D. andKnox, J.P. (2005). Arabinogalactan-proteinsare required for apical cell extension in themoss Physcomitrella patens. The Plant Cell,17: 3051-3065.

Peter H. RavenGeorge Engelmann Professor of BotanyDirector of the Missouri Botanical Gardenwww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONPeter H. Raven, a world leader in conservationand biodiversity, has devoted morethan 35 years at the Missouri BotanicalGarden to building a world-class institutionof research, education, and horticulturaldisplay. In working out systematic and evolutionaryproblems, he has utilized a broadapproach including data from cytogenetics,breeding systems, ecological preferences,pollination biology, and biogeographic considerations,as well as the more traditionaltypes of information.Peter Raven is co-editor of The Flora ofChina, a joint Chinese-American internationalproject that is producing a contemporaryaccount of the roughly 31,000 species ofplants of China. He has written numerousbooks and publications, both popular andscientific, and is senior author of The Biologyof Plants, the internationally best-sellingtextbook on botany now in its seventh edition,and of Environment, a leading textbookon the environment in its fifth edition.SELECTED PUBLICATIONSRaven, P.H., Johnson, G.B., Losos, J.B., Mason,K.A., Singer, S.R. (2008). Biology, Eighth edition.McGraw-Hill. Xxviii + 1259 pp.Raven, P.H., Evert, R., Eichhorn, S.E. (1999).Biology of Plants, Seventh edition, Freeman,Worth. xv + 944 pp.Raven, P.H., Axelrod, D.I. (1974).Angiosperm biogeography and past continentalmovements. Annals of the MissouriBotanical Garden, 61: 539-673.Ehrlich, P.R., Raven, P.H. (1969).Differentiation of populations. Science, 165:1228-1232.Ehrlich, P.R., Raven, P.H. (1965). Butterfliesand plants: a study in Coevolution.Evolution, 18: 586-603.Textbook authored by Peter H.Raven. George B. Johnson,Jonathan B. Losos, Kenneth A.Mason, and Susan R. Singer.39

Eric J. RichardsProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONMost eukaryotic genomes have fivenucleotide bases, rather than the familiar listof four. The fifth base is a modified form ofcytosine formed by the addition of a methylgroup. The Richards lab studies the mechanismsthat regulate eukaryotic DNA methylation,using molecular genetic and biochemicalapproaches applied to the flowering plantArabidopsis thaliana. In addition, we areinterested in understanding the function ofDNA modification and the broader evolutionarysignificance of epigenetic variation.SELECTED PUBLICATIONSRangwala, S.H. and Richards, E.J. (2007).Differential epigenetic regulation within anArabidopsis retroposon family. Genetics, 176:151-160.Woo, H.R., Pontes, O., Pikaard, C.S. andRichards, E.J. (2007). VIM1, a methylcytosine-bindingprotein required for centromericheterochromatinization. Genes &Development, 21: 267-277.Richards, E.J. (2006). Inherited epigeneticvariation — revisiting soft inheritance.Nature Reviews Genetics, 7: 395-401.Kuo, H-F., Olsen, K.M. and Richards, E.J.(2006). Natural variation in a subtelomericregion of Arabidopsis: implications for thegenomic dynamics of a chromosome end.Genetics, 173: 401-417.Rangwala, R., Elumalai, R., Vanier, C.,Ozkan, H., Galbraith, D.W. and Richards, E.J.(2006). Meiotically stable natural epiallelesof Sadhu, a novel Arabidopsis retroposon.PLoS Genetics, 2: e36.40The methylcytosine-binding protein VIM1 (darkgreen) functions to maintain centromeric heterochromatin.The centromere decondensationphenotype of vim1 mutants is illustrated on thetop left; blue, centromeric DNA; red, centromerespecifichistone H3 (HTR12). VIM1 bindingsmethylcytosine (m—C) and is proposed to actas an ubiquitin (Ub) ligase to modify centromerechromatin structure through direct modificationof nucleosomal histones or through modificationof one or more chromatin effector proteins (X).An alteration in centromeric chromatin structureleads to DNA hypomethylation and decondensationof the centromere repeats. Modified from Wooet al. Genes Dev (2007).

Barbara A. SchaalSpencer T. Olin Professorwww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONWork in Barbara Schaal’s lab focuses onthe evolutionary genetics of plants, often incollaboration with staff and students of theMissouri Botanical Garden. Research projectsspan the range from molecular evolution ofspecific DNA sequences to plant populationgenetics, systematics, and quantitative genetics.Schaal studies plant species native to theUnited States, tropical crops and their wildrelatives, and Arabidopsis.Rice growing in the Jeanette Goldfarb PlantGrowth FacilitySELECTED PUBLICATIONSBeck, J., Al-Shehbaz, I., O’Kane, S., Schaal,B. (2006). Further insights into the phylogenyof Arabidopsis (Brassicaceae) from nuclearAtmyb2 flanking sequence. MolecularPhylogenetics and Evolution, 42: 122-130.Miller, A. and Schaal, B. (2006).Domestication and the distribution of geneticvariation in wild and domesticated populationsof the Mesoamerican fruit tree Spondiaspurpurea. Molecular Ecology, 15: 1467-1480.Londo, J., Chiang, Y-C., Hung, K-H., Chiang,T-Y., and Schaal, B. (2006). Phylogeographyof Asian wild rice, Oryza rufipogon revealsmultiple independent domestications of cultivatedrice, Oryza sativa. Proceedings of theNational Academy of Sciences, 103: 9578-9583.Chung, K-F., Peng, C-I., Downie, S., Spalik,K. and Schaal, B. (2005). Molecular systematicsof the trans-Pacific alpine genusOreomyrrhis (Apiaceae): phylogenetic affinitiesand biogeographic implications. AmericanJournal of Botany, 92: 2054-2071.Schaal, B. and Leverich, W.J. (2005).Conservation Genetics: Theory and Practice.Annals of the Missouri Botanical Garden, 92:1-11.Caicedo, A. and Schaal, B. (2005).Heterogeneous evolutionary processes affectR gene diversity in natural plant populationsof Solanum pimpinellifolium. Proceedings ofthe National Academy of Sciences, 101:17444-17449.41

Paul S.G. SteinProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONTactile inputs excite neuronal circuits in thespinal cord that produce patterns of motoroutput responsible for specific rhythmicbehaviors. Paul S.G. Stein studies the physiologyof the interneuronal circuitry that generatesscratching in the turtle hindlimb(Earhart and Stein, 2000). He tests hypothesesof modular organization of the motorpattern generators in the spinal cord (Steinand Daniels-McQueen, 2002, 2003, 2004;Stein, 2005). He studies normal motor patternswith rhythmic alternation betweenagonists and antagonists. He also examinesmotor pattern variations, termed deletions,with rhythmic agonist activity and antagonistquiescence. He uses extracellular microsuctionelectrodes to obtain single-unit axonalrecordings of interneurons. These recordingsprovide support for the concept of modularspinal cord organization.SELECTED PUBLICATIONSStein, P.S.G. (2005). Neuronal control ofturtle hindlimb motor rhythms. Journal ofComparative Physiology A, Neuroethology,Sensory, Neural, and Behavioral Physiology,191: 213-229.Stein, P.S.G., Daniels-McQueen, S. (2004).Variations in motor patterns during fictiverostral scratching in the turtle: knee-relateddeletions. Journal of Neurophysiology, 91:2380-2384.Stein, P.S.G., Daniels-McQueen, S. (2003).Timing of knee-related spinal neurons duringfictive rostral scratching in the turtle. Journalof Neurophysiology, 90: 3585-3593.Stein, P.S.G., Daniels-McQueen, S. (2002).Modular organization of turtle spinalinterneurons during normal and deletionfictive rostral scratching. Journal ofNeuroscience, 22: 6800-6809.Earhart, G.M., Stein, P.S.G. (2000). Step,swim, and scratch motor patterns in theturtle. Journal of Neurophysiology, 84:2181-2190.Rhythms of motor neuron activities produced bythe turtle spinal cord in response to tactile stimulationof the turtle shell. Durations of agonistactivity are related to durations of antagonistquiescence. Figure 1A from P.S.G. Stein and S.Daniels-McQueen (2004), J. Neurophysiol. 91:2380-2384, used with permission of theAmerican Physiological Society.42

Alan R. TempletonCharles Rebstock Professor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONAlan Templeton’s work involves the applicationof molecular genetic techniques andstatistical population genetics to a varietyof evolutionary problems, both basic andapplied. He applies evolutionary approachesto clinical genetics, including the study ofthe genetics of coronary artery disease andthe evolution of the HIV virus within infectedpatients. He also applies evolutionarygenetics to conservation biology, with hismain current focus being the impact ofmanaged forest fires at the landscape levelupon the genetic population structure ofspecies inhabiting that landscape. Finally,he is interested in basic questions aboutevolution, such as the meaning of “species”and the mechanisms by which new speciesevolve, and recent human evolution.SELECTED PUBLICATIONSTempleton, A.R. (2005). Haplotype trees andmodern human origins. Yearbook of PhysicalAnthropology, 48: 33-59.Templeton, A.R., Maxwell, T., Posada, D.,Stengard, J.H., Boerwinkle, E. and Sing, C.F.(2005). Tree scanning: a method for usinghaplotype trees in genotype/phenotype associationstudies. Genetics, 169: 441-453.Templeton, A.R., Reichert, R.A., Weisstein,A.E., Yu, X.F. and Markham, R.B. (2004).Selection in context: patterns of naturalselection in the glycoprotein 120 region ofhuman immunodeficiency virus 1 withininfected individuals. Genetics, 167: 1547-1561.Brisson, J. A., Strasburg, J.L. and Templeton,A.R. (2003). Impact of fire management onthe ecology of collared lizard (Crotaphytuscollaris) populations living on the OzarkPlateau. Animal Conservation, 6: 247-254.A male collared lizard showing anovel yellow color on an Ozarkmountain recently colonized afterprescribed forest fires.43

Robert E. ThachDean, Graduate School of Arts & SciencesProfessor of Biologywww.biology.wustl.edu/facultyRESEARCH DESCRIPTIONOur chief area of study has in recent yearsbecome the ecology and epidemiology of vector-bornediseases. Of particular interest areErlichiosis, Southern Tick-associated Rash(Lyme-like) Illness, Tularemia, and RockyMountain Spotted Fever. The pathogensresponsible for these diseases are increasingin Missouri (long-recognized as a major centerof tick infestation in the United States),and pose a significant public health hazard.We utilize a combination of field and laboratoryapproaches to identify the animal reservoirswhich perpetuate these pathogens innature. Using this information we hope toshed light on why pathogens are not evenlydistributed geographically, but are oftenclustered in “hot spots.”SELECTED PUBLICATIONSAllan, B.F., Keesing, F. and Ostfeld, R.S.(February 2003). Effect of ForestFragmentation on Lyme Disease Risk.Conservation Biology, 17(1): 267-272.Butler, R.S., Arens, M., Harid, S.P., Paddock,C.D., Sumner, J.W., Rikhisa, Y., Unver, A.,Gaudreault-Keener, M., Manian, F.A., Liddell,A.M., Schmulewitz, N. and Storch, G.A. (July1999). New England Journal of Medicine,341(3): 148-155.Close collaborators in this study are GregoryA. Storch, Chief of Infectious Disease atSt. Louis Children’s Hospital, and Brian F.Allan, graduate student in the WashingtonUniversity Biology Department.Lone Star (Amblyomma americanum)ticks; left to right, adultmale, nymph, adult female44

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