Medical College of Wisconsin
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CURRICULUM VITAE

Joseph C. Besharse PhD, MA
Emeritus Professor
Department of Cell Biology Neurobiology and Anatomy
Marjorie & Joseph Heil Professor in Ophthalmology
OFFICE ADDRESS:
8701 Watertown Plank Road
Milwaukee, WI 53226
Phone: +1 (414) 9558063
Email: jbeshars@mcw.edu

EDUCATION:
1962 - 1965 Lyon College, Batesville, AR
1965 - 1966 BA, Hendrix College, Conway, AR
1966 - 1969 MA, Southern Illinois University at Carbondale, Carbondale, IL
1969 - 1973 PhD, Southern Illinois University at Carbondale, Carbondale, IL

POSTGRADUATE TRAINING AND FELLOWSHIP APPOINTMENTS:
1975 - 1977 Postdoctoral Fellow (NRSA), Columbia University College of Physicians and Surgeons, New York, NY

FACULTY APPOINTMENTS:
1972 - 1975 Assistant Professor, Biology, Old Dominion University, Norfolk, VA
1975 - 1977 NIH Postdoctoral Fellow, Ophthalmology and Cell Biology, Columbia University College of Physicians and Surgeons, NY, NY
1977 - 1980 Assistant Professor, Anatomy, Emory University School of Medicine, Atlanta, GA
1980 - 1984 Assistant Professor (Secondary Appt), Ophthalmology, Emory University School of Medicine, Atlanta, GA
1980 - 1984 Associate Professor (Tenured), Anatomy, Emory University School of Medicine, Atlanta, GA
1984 - 1989 Professor (Tenured), Anatomy and Cell Biology, Emory University School of Medicine, Atlanta, GA
1984 - 1989 Professor (Secondary Appointment), Ophthalmology, Emory University School of Medicine, Atlanta, GA
1985 - 1989 Director, Interdepartmental Graduate Cell Biology, Emory University, Atlanta, GA
1988 - 1989 Director, Graduate Program in Cell Biology, Division of Biological and Biomedical Sciences, Emory University, Atlanta, GA
1989 - 1997 Professor and Chairman, Anatomy and Cell Biology, The University of Kansas School of Medicine, Kansas City, KS
1989 - 1997 Professor (Secondary), Ophthalmology, The University of Kansas School of Medicine., Kansas City, KS
1997 - 2016 Professor and Chairman, Cell Biology, Neurobiology and Anatomy, Medical College of Wisconsin, Milwaukee, WI
1998 - 2016 Co-Director, Institutional Electron Microscopy Facility, Medical College of Wisconsin, Milwaukee, WI
2006 - 2017 Marvin Wagner Professor, Cell Biology, Neurobiology and Anatomy, Medical College of Wisconsin, Milwaukee, WI
2010 - 2014 Director, Interdisciplinary Graduate Program in Biomedical Sciences, Medical College of Wisconsin, Milwaukee, WI
2015 Marjorie & Joseph Heil Professor in Ophthalmology, Ophthalmology and Visual Sciences, Medical College of Wisconsin, Milwaukee, WI
2015 Director of Research, Ophthalmology and Visual Sciences, The Eye Institute, Medical College of Wisconsin, Milwaukee, WI
2016 Professor (Secondary Appointment), Cell Biology, Neurobiology and Anatomy, Medical College of Wisconsin, Milwaukee, WI

AWARDS AND HONORS:
1968 Preceptorships (for good teaching) , Southern Illinois University at Carbondale
1968 University Fellowship, Southern Illinois University at Carbondale
1971 University Fellowship , Southern Illinois University at Carbondale
1972 Preceptorship (for good teaching, Southern Illinois University at Carbondale
1975 Richard R. Kudo Award for and outstanding dissertation. , Southern Illinois University at Carbondale
1976 NIH Postdoctoral Fellowship (Individual NRSA), National Institutes of Health
1979 Research Career Development Award (5 years), National Institutes of Health
1993 Alcon Research Institute Award for outstanding research in Vision Science, Alcon Research Insitute
1996 BRAIN Scholar in Residence. , University of Houston
1998 Alumni Achievement Award, Southern Illinois University at Carbondale
1998 The Sydney Futterman Memorial Lecture, University of Washington, Seattle
2003 Appreciation for Contributions to Circadian Biology., Morehouse School of Medicine and Life Sciences Center
2004 Distinguished Service Award , The Association for Research in Vision and Ophthalmology
2006 Marvin Wagner Endowed Chair, Medical College of Wisconsin
2008 H. Talmage Dobbs Lectureship in Ophthalmology, Emory University School of Medicine
2008 Board of Trustees Medical College of Wisconsin, Medical College of Wisconsin
2009 Fellow, Association for Research in Vision and Ophthalmology
2010 Andre A. Belazs Prize for outstanding achievements in eye research. , International Society for Eye Research
2014 Fellow, American Association of Anatomy
2015 Distinguished Service Award, Medical College of Wisconsin
2015 Marjorie and Joseph Heil Professor of Ophthalmology, Medical College of Wisconsin
2016 Proctor Medal, Association for Research in Vision and Ophthalmology

MEMBERSHIPS IN HONORARY AND PROFESSIONAL SOCIETIES:
1971 - Present American Association for the Advancement of Science (Member)
1976 - Present Association for Research in Vision and Ophthalmology (Fellow, Past Vice President)
1977 - Present American Society for Cell Biology (Member, Past Council Member)
1978 - 2016 Society for Neuroscience (Member)
1983 - Present Society for the Study of Biological Rhythms (Member)
1989 - 2016 Association of Anatomy, Cell Biology and Neurobiology Chairs (Past President)
1989 - Present American Association of Anatomy (Fellow, Past Vice President)
1989 - Present International Society for Eye Research

EDITORSHIPS/EDITORIAL BOARDS/JOURNAL REVIEWS:
Editorship
1998 - 2000 Experimental Eye Research (Retina Section)
2008 - 2010 Encyclopedia of the Eye, Elsevier
2010 - 2011 Retina and its Disorders, Elsevier
Editorial Board
1985 - 1997 Experimental Eye Research
1988 - 1992 Investigative Ophthalmology and Visual Sciences
1990 - 1992 Visual Neuroscience
1997 - 2019 Molecular Vision
2010 - 2019 Eye and Brain

NATIONAL ELECTED/APPOINTED LEADERSHIP AND COMMITTEE POSITIONS:
1981 - 1982 Elected Chair, Program Committee, Division of Anatomy and Pathology, Association for Research in Vision and Ophthalmology and American Association of Anatomy
1981 - 1986 Study Section Member, Vision Research Review Committee, NEI/National Institutes of Health
1985 Chair, Local Arrangements Committee, Annual Meeting, Atlanta, GA, American Society for Cell Biology
1986 - 1989 Elected Member, Council, American Society for Cell Biology
1986 Invited Participant, Workshop on "Retinal Detachment", Bethesda, MD, National Institutes of Health
1990 Member, Public Affairs Committee, Association for Research in Vision and Ophthalmology
1992 - 1996 Study Section Member, Visual Sciences C Study Section, National Institutes of Health
1992 Organizer with Joe G. Hollyfield, Workshop on "The Role of the Cilium in Usher's Syndrome", Baltimore, MD, Foundation Fighting Blindness
1994 - 1996 Elected Vice President, Board of Directors, American Association of Anatomists
1994 - 1997 Elected Member, Council, Association of Anatomy, Neurobiology and Cell Biology Chairs
1996 - 2000 Chair, Public Affairs Committee, American Association of Anatomists
1999 - 2004 Elected Member, Board of Trustees, Association for Research in Vision and Ophthalmology
1999 Member, Department Chairs, National Caucus of Basic Biomedical Science
1999 - 2001 Elected President, Chair's Organization, Association of Anatomy, Neurobiology and Cell Biology Chairs
2000 - 2008 Member, Cell and Molecular Mechanisms of Disease Committee, Foundation Fighting Blindness
2001 Chair/Organizer, AANCB Chairs Presidential Symposium on “Advanced Model Systems for Early Vertebrate Development”, Orlando, FL, American Association of Anatomists
2004 Vice President, Board of Trustees, Association for Research in Vision and Ophthalmology
2006 - 2007 Elected Chair/Organizer, FASEB Conference on Biology and Chemistry of Vision, Federation of American Societies for Experimental Biology
2006 - 2009 Member, Fellow Selection Committee, American Association of Anatomists
2008 - 2009 Chair, Fellow Selection Committee, American Association of Anatomists

INTERNATIONAL ELECTED/APPOINTED LEADERSHIP AND COMMITTEE POSITIONS:
2012 Member, Paul Kayser International Award in Retina Research Review Committee, International Society for Eye Research Member
2014 Cell Biology Section Organizer, International Society for Eye Research Meeting, San Francisco, Program Committee
2016 Chair/Organizer, Program Committee Meeting, Tokyo Chair/Organizer, International Society for Eye Research
2016 Member, Belazs Prize Selection Committee, International Society of Eye Research

RESEARCH GRANTS/AWARDS/CONTRACTS/PROJECTS:
Active
Peer Review
Title:
RPE Signaling in Ocular Health and Disease
Source:
NIH/NEI
Role & Effort:
Multi-PI
PI:
Besharse and Link
Dates:
2018 - 2022
Direct Funds:
$1,814,304
  
Prior
Peer Review
Title:
Studies on Retinal Growth
Source:
Old Dominion University Research Foundation
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1973 - 1975
Direct Funds:
$4,400
  
Title:
Studies on a new cave salamander from West Virginia
Source:
American Philosophical Society
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1975
Direct Funds:
$400
  
Title:
Photoreceptor development in vitro
Source:
Emory University McCandless Fund
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1977
Direct Funds:
$1,000
  
Title:
Photosensitive membrane in developing ascidian photoreceptors.
Source:
National Science Foundation
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1978 - 1980
Direct Funds:
$2,045 ( Summer research at the Bermuda Biological Station)
  
Title:
Studies on disk renewal in retinal visual cells, R01-EY02414-01-33
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1978 - 1981
Direct Funds:
$125,360 (Original Award)
  
Title:
The Visual Cell - Pigment Cell Complex and Outer Segment Turnover
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1979 - 1984
  
Title:
Visual Cell - Pigment Cell Interface and Disc Turnover, R01-EY03222-01-05
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1979 - 1984
Direct Funds:
$286,791 (Original Award)
  
Title:
Studies on disk renewal in retinal visual cells, R01-EY02414-01-33
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1981 - 1986
Direct Funds:
$374,990 (Competitive Renewal)
  
Title:
Cyclic Nucleotides in the Retinal Pigment Epithelium
Source:
Fight-for-Sight, Inc. of New York City
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1982 - 1983
Direct Funds:
$3,000
  
Title:
Methoxyindoles in Retina: Function and Regulation
Source:
National Institutes of Health
Role & Effort:
Co-Investigator (15% Effort)
PI:
P. Michael Iuvone
Dates:
1983 - 1986
Direct Funds:
$276,933 (Original Award)
  
Title:
Electron Microscopy of Excitable and Secretory Cells, (JEOL 100cx)
Source:
National Institutes of Health
Role & Effort:
Program Director
PI:
J. C. Besharse
Dates:
1983 - 1984
Direct Funds:
$75,000 (NIH Instrumentation Grant: 50% funding of a transmission electron microscope)
  
Title:
Visual Cell - Pigment Cell Interface and Disc Turnover, R01-EY03222-01-35
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1984 - 1989
Direct Funds:
$510,189 (Competitive Renewal)
  
Title:
Studies on disk renewal in retinal visual cells, R01-EY02414-01-33
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1986 - 1991
Direct Funds:
$1,061,143 (Competitive Renewal)
  
Title:
Methoxyindoles in Retina: Function and Regulation
Source:
National Institutes of Health
Role & Effort:
Co-Investigator (15% effort)
PI:
P Michael iuvone
Dates:
1986 - 1989
  
Title:
Studies on Disc Renewal in Retinal Visual Cells, R01-EY02414 (Supplement)
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1989 - 1991
Direct Funds:
$71,000 (Equipment supplement for EY02414)
  
Title:
Visual Cell - Pigment Cell Interface and Disc Turnover, R01-EY03222-01-35
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1989 - 1994
Direct Funds:
$779,610 (Competitive Renewal)
  
Title:
Research Award
Source:
Alcon Research Institute
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1993 - 1997
Direct Funds:
$100,000 (Unrestricted for Research)
  
Title:
Visual Cell - Pigment Cell Interface and Disc Turnover, R01-EY03222-01-35
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1994 - 1999
Direct Funds:
$1,597,973 (Competitive Renewal)
  
Title:
Circadian Rhythms in Zebrafish, F49620-94-1-0314
Source:
United States Air Force
Role & Effort:
PI
PI:
J. C. Besharse (with Gregory Cahill)
Dates:
1994 - 1996
Direct Funds:
$60,000
  
Title:
Studies on disk renewal in retinal visual cells, R01-EY02414-01-33
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
1996 - 2001
Direct Funds:
$1,302,853 (Competitive Renewal)
  
Title:
Visual Cell - Pigment Cell Interface and Disc Turnover, R01-EY03222-01-35
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
2000 - 2007
Direct Funds:
$1,766,940 (Competitive Renewal)
  
Title:
Studies on disk renewal in retinal visual cells, R01-EY02414-01-33
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
2001 - 2005
Direct Funds:
$1,420,250 (Competitive Renewal)
  
Title:
Spectral confocal Microscope, 1S10RR015679-01
Source:
National Institutes of Health
Role & Effort:
Program Director
PI:
J. C. Besharse
Dates:
2001 - 2002
Direct Funds:
$351,000 (Large Instrumentation Grant)
  
Title:
T32-EY014537: Research Training Program in Vision Science
Source:
National Institutes of Health
Role & Effort:
Program Director
PI:
J. C. Besharse
Dates:
2002 - 2008
Direct Funds:
$838,970 (Original 5 yr award )
  
Title:
Studies on disk renewal in retinal visual cells, R01-EY02414-01-33
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
2005 - 2011
Direct Funds:
$1,050,915 (Competitive Renewal)
  
Title:
FASEB Summer Research conference on the “Biology and Chemistry of Vision”, 1 R13 EY018289
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
2006 - 2007
Direct Funds:
$30,000 (Conference Grant)
  
Title:
Visual Cell - Pigment Cell Interface and Disc Turnover, R01-EY03222-01-35
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
2008 - 2011
Direct Funds:
$1,136,250 (Competitive Renewal)
  
Title:
T32-EY014537: Research Training Program in Vision Science
Source:
National Institutes of Health (NEI
Role & Effort:
Program Director
PI:
J. C. Besharse
Dates:
2008 - 2013
Direct Funds:
$941,445 (Competitive Renewal)
  
Title:
Technology Development and Animal Models for Analysis of Photoreceptor Turnover
Source:
Advancing Healthier Wisconsin
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
2009 - 2011
Direct Funds:
$200,000
  
Title:
Regulating Cerebral Microcirculation by Astrocytes, R01-HL033833, Yrs. 25-28
Source:
National Institutes of Health (NHLBI)
Role & Effort:
Co-Investigator (10% effort)
PI:
D.R. Harder
Dates:
2010 - 2012
  
Title:
Visual Cell - Pigment Cell Interface and Disc Turnover, R01-EY03222-31-35
Source:
National Institutes of Health
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
2011 - 2017
Direct Funds:
$2,200,000 (Competitive Renewal)
  
Title:
A Role for the Circadian De-Ubiquitinase, USP2, in Tumorigenesis
Source:
MCW Cancer Center
Role & Effort:
PI
PI:
J. C. Besharse
Dates:
2011 - 2013
Direct Funds:
$50,000
  
Title:
T32-EY014537: Research Training Program in Vision Science
Source:
National Institutes of Health (NEI)
Role & Effort:
Program Director
PI:
Joseph C Besharse
Dates:
2013 - 2018
Direct Funds:
$1,489,015 (Second Competing Renewal)
  

INVITED LECTURES/WORKSHOPS/PRESENTATIONS:
International
Besharse (Invited), "Light and Photosensitive Membrane Turnover in Rod Photoreceptors in vitro", IV International Symposium on the Structure of the Eye, Guadalajara, Mexico, 08/1980
Besharse (Invited, "Methoxyindoles and Photoreceptor Metabolism: Activation of Disc Shedding,", Vth International Congress of Eye Research, Workshop on "Drug Induced Modification of Retinal Metabolism", Eindhoven, Netherlands, 10/1982
Besharse (Invited), "Temporal Regulation of Rhythmic Membrane Turnover: A Model for Control of Light Evoked and Circadian Events,", VIIth International Congress of Eye Research, Symposium on "Cyclic Retinal Processes", Nagoya, Japan, 10/1986
Besharse (Invited), "Regulation of Rhythmic Photoreceptor Metabolism: A Role for Dopamine and Melatonin", International Symposium on "Retinal Cell Biology", Royal Ontario Museum, Canada, 04/1987
Besharse (Invited), "Retinal Rhythmicity: Melatonin as a Local Modulator and Circadian Signal", Canadian Federation of Biological Societies, Symposium on "The Retina as a Model for Neurobiology", Calgary, AB, Canada, 06/1989
Besharse (Symposium organizer and Invited Speaker), "Anchorage of Unique Cell-Surface Glycoconjugates of the Connecting Cilium to Microtubules of the Axoneme: and "The Circadian Clock in the Eye: Ocular Melatonin Metabolism", IXth International Congress of Eye Research, Helsinki, Finland, 08/1990
Besharse (Invited), "The Clock in the Retina: Regulation of Retinal Rhythmicity by Light and Dopamine", International Society of Neurochemistry, Symposium on "Signal Transmission in the Retina, Sydney, Australia, 07/1991
Besharse (Invited), "The Clock in the Retina: Role of Photoreceptors", Xth International Congress of Eye Research, Symposium on Retinal Photoreceptors, Stresa, Italy, 09/1992
Besharse (Invited), “The Photoreceptor Circadian Clock: Regulation of Melatonin Metabolism and Gene Transcription”, Les Seminaires Ophthalmologiques d’Ipsen, Development and Differentiation of the Retina, Royaumont, France, 03/1995
Besharse (Invited), “The Photoreceptor Circadian Clock: Physiological and Pathological Implications”, International Symposium on Retinal Degeneration, Sendai, Japan, 10/1996
Besharse (Invited), “Gene Regulation and the Photoreceptor Circadian Clock”, 7th Sapporo Symposium on “Biological Rhythms”, Sapporo, Japan, 09/1997
Besharse (Invited), “Photoreceptor Intersegmental Transport and Retinal Degeneration, A conserved pathway common to motile and sensory cilia”, International Symposium on Retinal Degeneration, Burgenstock, Switzerland, 10/2002
Besharse (Invited), "Cellular/Molecular Organization of circadian Oscillations in the Retina", Gordon Research Conference on "Chronobiology", Il Ciccio, Barga, Italy, 05/2003
Besharse (Invited), "Regulation of Photoreceptor Per1 and Per2 by Light, Dopamine and a Circadian Clock", International Congress of Eye Research, Sydney, Australia, 08/2004
Besharse (Invited), Belazs Prize Lecture: "The Intersection of Circadian and Cellular Processes Regulating"Photoreceptor Membrane Turnover", International Society of Eye Research XIXth Biennial Meeting, Montreal, QC, Canada, 07/2010
Besharse (Invited), Plenary Lecture: "Intraflagellar transport and the regulation of photoreceptor outer segment assembly," Plenary Lecture I, ARVO "Ocular Cell Biology" Research Conference, Vancouver, BC, Canada, 09/2011
Besharse and Lewis (Session Organizer and Speaker; unable to attend. Talk given by Tylor Lewis), Microtubule Organization and the Early Development of Photoreceptors Outer Segments., International Society for Eye Research (ISER). XXII Meeting, Tokyo, Japan, 09/2017
 
National
Besharse, "Morphology of the Eyes of Some Troglobitic Urodeles", Symposium on Cave Biology, Western Kentucky University, Bowling Green, KY, 04/13/1973
Besharse (Invited), "Retinal Degeneration in the Ozark Cave Salamander", Retinitis Pigmentosa Foundation Workshop, Endicott House, Dedham, MA, 05/1976
Besharse (Invited), "Light and Membrane Biogenesis in Rod Photoreceptors of Vertebrates", Symposium on "Effects of Constant Light on Visual Processes", Florida State University, Tallahassee, FL, 04/1979
Besharse (Invited), "Light Evoked Disc Shedding by Rod Photoreceptors in vitro", Vision in Charleston, II, Medical University of South Carolina, Charleston, SC, 05/1980
Besharse (Invited), "Precursor Delivery to Photoreceptor Outer Segments During Periods of Rapid Disc Assembly,", IV International Congress for Eye Research, Symposium on "Control of Photoreceptor Renewal", New York, NY, 09/1980
Besharse and Iuvone (Invited), "Melatonin Biosynthesis in Retina and its Possible Roles in Photoreceptor Physiology,", 67th Annual Meeting of the Federation of American Societies for Experimental Biology, Symposium on "Catecholamines and Indoleamines in Retina", Chicago, IL, 04/1983
Besharse (Invited), "Photoreceptor Disc Shedding: Mechanism and Regulation", Symposium on "Cell Biology of the Retina", Baylor College of Medicine, Houston, TX, 11/1983
Besharse (Invited), "Photoreceptor Disc Shedding and Phagocytosis: Mechanism and Regulation,", American Society for Cell Biology, Symposium on Control of Photoreceptor Renewal", Kansas City, KS, 11/1984
Besharse (Invited), "Melatonin and Dopamine Interaction in the Control Retinal Photoreceptor Rhythmicity", Gordon Research Conference on "Chronobiology", Plymouth State College, Plymouth, NH, 07/1985
Besharse (Invited), "Role of Postreceptoral Neurons in Control of Rhythmic Behavior of Photoreceptors,", FASEB Summer Research Conference on "Biology and Chemistry of the Visual Process", Saxtons River, VT, 08/1985
Besharse (Symposium Organizer and Speaker), "Cell Biology of Retina", Department of Ophthalmology, Emory University, Atlanta, GA, 10/1985
Besharse and Yazula (Symposium Organizers), Retinal Organization and Circuitry, International Congress of Eye Research, San Francisco, CA, 1986
Besharse (Invited), Dopaminergic Regulation of Light-Adaptive Responses in Xenopus Photoreceptors", FASEB Summer Research Conference on "Biology and Chemistry of Vision", Copper Mountain, CO, 07/1987
Besharse (Invited), "A Role for Melatonin and Dopamine in Circadian Regulation in the Vertebrate Retina", Gordon Research Conference on "Chronobiology", Plymouth State College, Plymouth, NH, 07/1987
Besharse (Organizer and Speaker), "Cell Biology of Photoreceptors and Pigment Epithelium", American Society for Cell Biology, Retinal Cell Biology Special Interest Subgroup Meeting, St. Louis, MO, 1987
Besharse (Invited), "Dopaminergic Regulation of Light-Adaptive Responses in Retina", Winter Conference on "Brain Research", Steamboat Springs, CO, 02/1988
Besharse (Invited), "Regulation of Retinal Melatonin Secretion", Symposium of Society for Research on Biological Rhythms, Charleston, SC, 05/1988
Besharse (Invited), "Membrane Assembly in Vertebrate Photoreceptors: Mechanism of Opsin Delivery", 2nd International Congress of Comparative Physiology and Biochemistry, Baton Rouge, LA, 08/1988
Besharse (Invited), "Dopamine as a Paracrine Effector in the Regulation of Cone Photoreceptor Motility", VIIIth International Congress of Eye Research, San Francisco, CA, 09/1988
Besharse (Invited), "Identification and Characterization of Glycoconjugates Unique to the Photoreceptor Connecting Cilium.", FASEB Summer Research Conference on "Biology and Chemistry of Vision", Saxtons River, VT, 06/1989
Besharse (Invited), "The Clock in the Eye: Regulation of Retinal Rhythmicity by Melatonin and Dopamine", Symposium on Retinal Cell Biology, Baylor College of Medicine, Houston, TX, 11/1989
Besharse (Invited), "The Clock in the Retina: Regulation of Retinal Rhythmicity by Dopamine and Melatonin.", Frederick E. Sampson Symposium on Neurochemistry, University of Kansas Medical Center, 10/1990
Besharse (Invited), "The Clock in the Eye: Regulation of Retinal Rhythmicity by Melatonin and Dopamine", The Broadhurst Foundation Visiting Lecture Series, Eye Research Institute, Harvard University, 04/1991
Besharse (Invited), "In Vitro Analysis of a Retinal Circadian Clock", FASEB Summer Research Conference on the "Biology and Chemistry of Vision", Copper Mountain, CO, 07/1991
Besharse (Invited), "In Vitro Analysis of a Retinal Circadian Clock", Advanced Concepts in Vision Science: Circadian Rhythms, Department of Ophthalmology, Harvard Medical School, Boston, MA, 12/1991
Besharse (Workshop Organizer with Joe G Hollyfield and Speaker), "Toward a Molecular Structure of the Photoreceptor Cilium", National Retinitis Pigmentosa Foundation Fighting Blindness, Workshop on "The role of the Cilium in Usher's Syndrome, Baltimore, MD, 07/1992
Besharse (Invited), "The Photoreceptor Circadian Clock: Melatonin, Dopamine and Rhythmic Regulation", The Helen Keller Eye Research Foundation Seminar Series, University of Alabama Medical Center, Birmingham, AL, 05/1993
Besharse (Invited), “Phagocytosis of Photosensitive Membranes by Retinal Pigment Epithelium In Situ”, Experimental Biology 1994, Symposium on “The Retinal Pigment Epithelium In Vitro: A Dynamic View of a Multifunctional Cell”, Anaheim, CA, 04/1994
Besharse (Invited), Award Lecture: “The Clock in the Eye: Localization in Photoreceptors and Physiological Relevance”, The Alcon Research Institute Annual Awards Program, Fort Worth, TX, 04/1994
Besharse (Chair and Organizer), “Disc Morphogenesis in Photoreceptors: The Final Steps”, ARVO, Special Interest Subgroup Meeting, Sarasota, FL, 05/1994
Besharse (Invited), “The Photoreceptor Circadian Clock: Physiological Significance and Role in Transcription Regulation”, Advances in Vision Neuroscience, A Symposium honoring Professor Adoplh Cohen, Washington University School of Medicine, St. Louis, MO, 09/1994
Besharse (Organizer and Speaker), “Cellular and Molecular Biology of Circadian Clocks”, American Association for Cell Biology, Special Interest Subgroup Meeting, San Francisco, CA, 12/1994
Besharse (Invited), “The Photoreceptor Connecting Cilium and Disk Morphogenesis”, FASEB Summer Research Conference on the "Biology and Chemistry of Vision", Copper Mountain, CO, 07/1995
Besharse (Invited), The Photoreceptor Circadian Clock, Melatonin Synthesis and Gene Regulation, Winter Brain Research Conference, Snowmass, CO, 02/1996
Besharse (Organizer with Stuart Mangel and Speaker), “The Photoreceptor Circadian Clock and Regulation of Gene Expression”, Association for Research in Vision and Ophthalmology, Minisymposium on “Circadian Rhythmicity in the Retina: Cellular, Molecular and Functional Aspects”, Ft. Lauderdale, FL, 04/1996
Besharse (Invited), “The Photoreceptor Circadian Clock and Regulation of Gene Expression”, U.S. Army Research Office Conference on “Response and Adaptation to the Environment”, Research Triangle Park, NC, 10/1996
Besharse (invited), “The Photoreceptor Circadian Clock and Regulation of Gene Expression”, BRAIN Scholar in Residence, Program in “Basic Research and Academic Initiatives in Neuroscience”, University of Houston, Houston, TX, 11/1996
Besharse (Invited), “Circadian Gene Regulation in Photoreceptors”, Keystone Symposium on “Ocular Cell and Molecular Biology”, Tamarron, CO, 01/1997
Besharse (Invited Session Organizer and Speaker), “The photoreceptor circadian clock in vertebrates: regulation of gene transcription”, FASEB Summer Research Conference on the "Biology and Chemistry of Vision", Keystone, CO, 07/1997
Besharse (Invited Session Organizer and Speaker), Retinal Rhythmicity, Pineal Gland Gordon Conference, Ventura, CA, 02/1998
Besharse (Invited), “The Ocular Circadian Clock”, The Sydney Futterman Memorial Lecture, Ophthalmology, University of Washington, Seattle, Seattle, WA, 04/1998
Besharse (invited), “‘End of Progress’ or ‘Endless Frontier’”, Graduation Address to College of Sciences and Recipient of Alumni Achievement Award, Southern Illinois University, Carbondale, IL, 05/1998
Besharse (invited), “Clock and Clock-Controlled Genes in Retinal Circadian Function”, Loris and David Rich Lecture Series in Visual Science, University Of Alabama at Birmingham, Birmingham, AL, 09/1998
Besharse (Invited), “Clock and Clock-Controlled Genes in Retinal Circadian Function”, Keystone Symposium on “Ocular Cell and Molecular Biology”, Keystone, CO, 02/1999
Besharse (Invited), “Light- and Circadian Clock-Controlled Genes in the Retina”, Winter Brain Research Conference, Keystone, CO, 02/2000
Besharse (Invited Symposium Organizer and Speaker), “The Retinal Circadian Clock: Regulation of Gene Expression and Physiology”, Association for Research in Vision and Ophthalmology, Symposium on “Circadian Clocks and their Impact on Retinal Function and Disease”, Ft. Lauderdale, FL, 04/2000
Besharse (Invited), “Intraflagellar transport: a model for intersegmental transport in vertebrate photoreceptors”, International Symposium on Retinal Degeneration, Durango, CO, 10/2000
Besharse (Invited), "Light Regulation of the Xenopus Period 2 gene in the Retina", Keystone Symposium on "Molecular Clocks: Regulation of Circadian Behavioral Rhythms", Tahoe City, CA, 03/2001
Besharse (Invited), "Phase Shifting in Photoreceptors Through Light and Dopamine -induced Transcription of Per2", FASEB Summer Research Conference on the "Biology and Chemistry of Vision", Tucson, AZ, 06/2001
Besharse (Invited), “Intersegmental Transport in Photoreceptors. Evidence for an Ancient, conserved Mechanism in Motile and Sensory Cilia”, Distinguished Lecture Series , Cole Eye Institute, The Cleveland Clinic, Cleveland, OH, 05/2002
Besharse (Invited), “Intraflagellar transport and the assembly photoreceptor outer segments”, FASEB Summer Research Conference on the "Biology and Chemistry of Vision", Tucson, AZ, 06/2003
Besharse (Invited), "Vertebrate Photoreceptors as Sensory Cilia: A Conserved Transport Mechanism in Motile and Sensory Cilia", Gordon Research Conference on "Ocular Development", Roger Williams University, Bristol, RI, 06/2004
Besharse (Symposium Organizer and Speaker), “Cellular Pathways and Neurodegeneration; Microtubule based ciliary transport and assembly of the photoreceptor outer segment.”, Experimental Biology 2005 Annual Meeting, Joint ARVO/AAA, (Symposium held a both EB2005 and ARVO 2005 Meetings), San Diego, CA and Ft. Lauderdale, FL, 04/2005
Besharse (Invited Session Organizer and Speaker), “Confocal and Multiphoton imaging of IFT57, 52, and 88”, Ninth Annual Vision Research Conference on Imaging, Ft. Lauderdale, FL, 05/2005
Besharse (Invited), "Microtubule based transport of photoreceptor specific cargo through the connecting cilium", ARVO Summer Research Conference on "Ocular Cell and Molecular Biology", Sarasota, FL, 09/2005
Besharse (Invited), "Genetic Analysis of Circadian clocks in the Retina", Jerome Sutin Memorial Symposium, A Celebration of Molecular Neuroscience, Emory University, Atlanta, GA, 04/2006
Besharse (Invited), “Rhodopsin and Intraflagellar Transport in Photoreceptors”, Tenth Annual Vision Research Conference on Rhodopsin, Ft. Lauderdale, FL, 05/2006
Besharse (Invited), “Motors and Cargo for Intraflagellar Transport in Photoreceptors”, Eleventh Annual Vision Research Conference on Neurodegeneration, Ft. Lauderdale, FL, 05/2007
Besharse (Conference Chair and Organizer), Biology and Chemistry of Vision, FASEB Summer Research Conference on "The Biology and Chemistry of Vision", Snowmass Village, CO, 06/2007
Besharse (Invited), “Motors and Cargo for Intraflagellar Transport in Photoreceptors”, FASEB Summer Research Conference on "Cilia and Flagella", Saxtons River, VT, 08/2007
Besharse (Invited), "Photoreceptor specific IFT: identification of Kif17 as an accessory IFT motor and Dnajb6 as an IFT linker for retinal guanylyl cyclase in photoreceptors", ARVO Summer Research Conference on "Ocular Cell and Molecular Biology", Sarasota, FL, 09/2007
Besharse (Invited), "Motors and Cargo for Intraflagellar Transport in Photoreceptors", H. Talmage Dobbs Lectureship in Ophthalmology, Emory Eye Center, Emory University School of Medicine, Atlanta, GA, 02/2008
Besharse (Invited), “Molecular Motors and Cargo of Photoreceptor IFT”, 2nd International Conference on “Ocular Cell Biology”, Paradise Point Resort, San Diego, CA, 09/2008
Besharse (Invited), “Regulation of Photoreceptor KIF17 Kinesin Transport into the Outer Segment by CamKII”, Association for Research in Vision and Ophthalmology, Symposium on “Photoreceptor Cilia and Associated Disease”, Ft. Lauderdale, FL, 04/2009
Besharse (Invited), “Phosphorylation of Kif17 Kinesin Regulates IFT Transport into the Outer Segment”, FASEB Summer Research Conference on the "Biology and Chemistry of Vision", Snowmass, CO, 06/2009
Besharse (Invited), “Photoreceptors and Ciliogenesis: KIF17 Regulates IFT Transport and Outer Segment Turnover,” Seminar, H. Talmage Dobbs Lecture. Emory University School of Medicine, Atlanta, GA, 07/2009
Besharse (Invited), “Photoreceptors and Ciliogenesis: KIF17 Regulates IFT Transport and Outer Segment Turnover,” Seminar, Department of Ophthalmology, Columbia University College of Physicians and Surgeons, New York, NY, 12/2009
Besharse (Invited), "Circadian Regulation of Photoreceptor Outer Segment Turnover", Gordon Research Conference on "The Pineal Gland", Galveston, TX, 02/2010
Besharse (Invited), “Regulation Circadian Photoreceptor Outer segment Turnover by Kinesin 17”, Jules Stein Eye Institute, UCLA Medical School, Los Angeles, CA, 11/2010
Besharse (Invited), "Kinesin Trafficking in the Cilium and Regulation of Photoreceptor Membrane Turnover", Pioneers in Neuroscience Seminar Series, University of Buffalo, Buffalo, NY, 10/2011
Besharse (Invited), “The role of two kinesin 2 family motors in photoreceptor ciliogenesis and turnover of the distal outer segment”, 14th Vision Research Conference: Retina Ciliopathies, from Genes to Mechanisms and Treatment, Ft. Lauderdale, FL, 05/2012
Besharse (Invited Cell Biology Section Organizer and Speaker), "Axoneme Organization and IFT in Zebrafish and Mammalian Photoreceptors", International Society of Eye Research XXIth Biennial Meeting, San Francisco, CA, 07/2014
Besharse, “Massive Recruitment of Subretinal, Ccr2+ Macrophages in Mfrp Deficient Mice Does Not Impact Retinal Degeneration," Symposium, Association for Research in Vision and Ophthalmology Symposium, Denver, CO, 05/2015
Besharse (Invited), Local Circadian Clocks at the Interface of Ocular Physiology, Proctor Medal Lecture, Association of Research in Vision and Opthalmology, Seattle, WA, 05/2016
Besharse (Invited), Kif17 Phosphorylation, ciliary distal “tipping” and cone photoreceptor outer segment turnover, FASEB Summer Research Conference on the "Biology and Chemistry of Vision", Steamboat Springs, CO, 06/2017
Besharse Invited, “The Molecular Clockwork in Eye: Organization and Significance”., Stephenson Distinguished Lecture Series In Ophthalmology., University of Louisville., 2018
 
Regional
Besharse (Invited), NOTE: Department Seminars NOT included. Only invited speaker events are listed for meetings), Many Colleges and Universities, 1973 - 2016
Besharse (Invited), “The Cell Biology of Photoreceptor Membrane Turnover: Kinesins, IFT and the Phototransduction Machinery”, Symposium: “Greater than the Sum: Advances in Correlative Microscopy”, Northwestern University, Evanston, IL, 04/2009
Besharse (Invited), Keynote Lecture: De-Ubiquitination within the Molecular Clockwork and Circadian Sensitivity to Light, Midwest Vision Research Symposium, Department of Ophthalmology, University of Iowa, Iowa City, IA., 07/2011
Besharse (Invited), Subretinal Macrophages and Photoreceptor Loss in MFRP Deficient Mice: A Cautionary Tale, 2016 Spring Vision Symposium, University of Wisconsin, Madison, Madison, WI, 04/2016
 

COMMITTEE SERVICE:
Medical College of Wisconsin
1977 - 1989 Member, Emory: Dissertation Committees, Multiple Students, Cell Biology and Anatomy, Emory Medical School
1978 - 1981 Member, Departmental Committee on Graduate Training, Cell Biology and Anatomy, Emory University
1979 Member, Departmental Faculty Search Committees, Cell Biology and Anatomy, Emory University
1979 - 1980 Member, Natural Science Divisional Review Committee, Emory University
1981 Member, Departmental Faculty Search Committees, Cell Biology and Anatomy, Emory University
1983 - 1989 Member, Ad-Hoc Subcommittee of the Committee on Appointments and Promotions for Medical School, Emory University
1983 Member, Departmental Faculty Search Committees, Cell Biology and Anatomy, Emory University
1984 - 1988 Member, Long-Range Planning Committee of Emory University School of Medicine, Emory University
1988 Chair, Cell Biology Program Committee, Graduate Division of Biological Sciences, Emory University
1990 Member, Search Committee for Chair of Preventive Medicine, University of Kansas Medical Center
1990 - 1992 Member, University Tenure and Promotions Committee, University of Kansas Medical Center
1990 Member, Search Committee for Chair of ENT, University of Kansas Medical Center
1990 - 1997 Departmental Teaching and Curriculum Review Committee, Anatomy and Cellular Biology, University of Kansas Medical Center
1991 Member, Committee for Departmental and Chairmanship Review, University of Kansas Medical Center
1991 - 1992 Member, Advisory Committee on Research Space, University of Kansas Medical Center
1991 Member, Flow Cytometry Facility Review Committee, University of Kansas Medical Center
1991 - 1994 Chair, Monoclonal Antibody Facility Review Committee, University of Kansas Medical Center
1991 - 1992 Member, Marion-Merrill Dow Endowed Professorship Search Committee, University of Kansas Medical Center
1992 Member, Robert Wood Johnson Steering Committee, University of Kansas Medical Center
1992 - 1993 Chair, Search Committee for Chairperson of the Department of Physiology, University of Kansas Medical Center
1992 - 1994 Chair, Scientific Advisory Committee of Alzheimer's Research Center, University of Kansas Medical Center
1993 Member, Search Committee for Executive Dean of the School of Medicine, University of Kansas Medical Center
1994 Member, Search Committee for Halls Endowed Professorship in Molecular Medicine, University of Kansas Medical Center
1994 - 1995 Member, School of Medicine Education Council, University of Kansas Medical Center
1994 Chair, Confocal Facility Planning Group, University of Kansas Medical Center
1994 Member, Search Committee for Associate Dean for Research, University of Kansas Medical Center
1995 Member, Search Committee for Director and Associate Director of PCPE, University of Kansas Medical Center
1995 - 1996 Chair, Transgenic Facility Planning Committee, University of Kansas Medical Center
1996 Chair, Personal Computer Allocation Committee, University of Kansas Medical Center
1997 Member, Genetics Initiative Committee, Medical College of Wisconsin
1997 Chair, Electron Microscopy Facilities Review Committee, Medical College of Wisconsin
1997 - 1999 Member, MSTP Training Program Advisory Committee, Medical College of Wisconsin
1999 Chair, Search Committee for Director of Human and Molecular Genetics Center, Medical College of Wisconsin
1999 - 2008 Member, Dean’s Research Space Advisory Committee, Medical College of Wisconsin
1999 - 2016 Member, Executive Committee of Medical Scientists Training Program, Medical College of Wisconsin
2000 - 2003 Member, Curriculum and Evaluation Committee, Medical College of Wisconsin
2001 - 2006 Co-Chair, Animal Research Center Oversight Committee, Medical College of Wisconsin
2003 Member, Dean’s Research Planning Committee, Medical College of Wisconsin
2004 Member, Laboratory Design Evaluation Committee (new Research Building), Medical College of Wisconsin
2004 - 2005 Chair Representative, Faculty Council, Medical College of Wisconsin
2006 - 2009 Chair, BRC Advisory Committee, Medical College of Wisconsin
2008 - 2010 Member, Board of Trustees, Medical College of Wisconsin
2009 Member, Biochemistry Chair Search Committee, Medical College of Wisconsin
2009 - 2010 Member, Dean’s Education and Innovation Committee, Medical College of Wisconsin
2009 - 2010 Member, Ophthalmology Research Search Committee, Medical College of Wisconsin
2011 Member, Curriculum Development Nominating Committee of the Faculty Council, Medical College of Wisconsin
2011 - Present Internal Advisory Committee, Cancer Center, Medical College of Wisconsin
2012 - Present Member, Mentoring Oversight Board (MOB), Medical College of Wisconsin
2012 - 2015 Member, Medical School Curriculum and Evaluation Committee, Medical College of Wisconsin
2013 - 2015 Chair, Search Committee for Senior Associate Dean for Research at MCW, Medical College of Wisconsin
2014 - 2016 Member, Research Council, Medical School
2016 Member, Distinguished Service Award Selection Committee, Medical College
 

MCW TEACHING ACTIVITIES:
Medical Student Education
1997 - 2013 Lectures/Discussion/Laboratories in Cell and Tissue Biology (2-5 sessions annually)
1999 - 2018 Mentored/Sponsored 3 Summer Medical Student Research Rotations
 
Graduate Student Education
1997 - 2017 Primary Mentor of 7 Graduate Students
1998 - 2012 Taught required course in "Analysis of Scientific Data" (10-12 sessions annually))
1998 - 2016 Course in Advanced Cell Biology (2-3 sessions biannually)
1999 - 2018 Membership on 8 PhD Advisory Committees
1999 - 2017 PhD Qualifying Exams (1-2 annually)
1999 - 2018 Lectures/Discussions in Required Course Molecular Cell Biology (5-6 sessions annually)
2003 - 2018 Course in Biology and Chemistry of Vision (2-3 sessions biannually)
 
Postdoctoral Fellow Education
1997 - 2018 Mentored 14 Post-Doctoral Trainees
 

EXTRAMURAL TEACHING:
Medical Student Education
1977 - 1989 Emory University Medical School, 6-10 Lectures/Laboratories annually
1989 - 1997 University of Kansas School of Medicine, Teaching 5-7 lectures/labs annually
 
Graduate Student Education
1977 - 1989 Emory University School of Medicine, Primary Mentor for 4 PhD Students
1977 - 1989 Emory University School of Medicine, Member of 10 Dissertation Committees
 
Postdoctoral Fellow Education
1977 - 1989 Emory University School of Medicine, Mentored 5 Post-Doctoral Trainees
1989 - 1997 University of Kansas School of Medicine, Mentored 7 Post-doctoral Trainees
 

MCW STUDENTS, FACULTY, RESIDENTS AND CLINICAL/RESEARCH FELLOWS MENTORED:
Graduate Students
MS Students Advised
2000 Minhong Zhuang, Differential Regulation of Two Period Genes in the Xenopus Eye, Medical College of Wisconsin
2003 Mei Li, Intraflagellar Transport Protein, IFT88, Directly Interacts with the Chaperone Protein MRJ, Medical College of Wisconsin
PhD Students Advised
2003 Sheila A. Baker, A Study of Intraflagellar Transport in Vertebrate Photoreceptors, Medical College of Wisconsin
2009 Christine Insinna Kettenhofen, Role of Kinesin 2 Motors in Photoreceptor Cilium Assembly and Maintenance, Medical College of Wisconsin
2011 Heather M. Dehlin, De-Ubiquitinase Modulated Circadian Clock Re-Setting by Light, Medical College of Wisconsin
2013 Joseph Fogerty, A Novel Mouse Mutant Resembling Age-related Macular Degeneration, Medical College of Wisconsin
2017 Tyler R Lewis, Microtubule Motor in the Ciliary Photoreceptor, Medical College of Wisconsin
 
Postdoctoral Researchers
1996 - 2000 Yunxia Wang, Research Associate/Research Assistant Professor, Medical College of Wisconsin
1998 - 2000 Pamela Megaw, Postdoctoral Fellow, Medical College of Wisconsin
2001 - 2004 Xinrong Lu, Postdoctoral Fellow, Medical College of Wisconsin
2001 - 2004 Jun Sun, Postdoctoral Fellow, Medical College of Wisconsin
2001 - 2006 Katherine Luby-Phelps, Research Associate/Assistant Professor, Medical College of Wisconsin
2002 - 2003 Qingjiong Zhang, Postdoctoral Fellow, Medical College of Wisconsin
2004 - 2007 Reshma Bhowmick, Postdoctoral Fellow, Medical College of Wisconsin
2005 - 2008 Geetha Yadav, Research Associate, Medical College of Wisconsin
2005 - 2006 Carl Strayer, Postdoctoral Fellow, Medical College of Wisconsin
2005 - 2010 David Lourim, Research Associate/Research Scientist, Medical College of Wisconsin
2009 - 2011 Kirsten Janisch, Postdoctoral Fellow, Medical College of Wisconsin
2010 - 2014 Jason Bader, Postdoctoral Fellow, Medical College of Wisconsin
2015 - Present Steven Henle, Postdoctoral Fellow, Medical College of Wisconsin
 

EXTRAMURAL STUDENTS, FACULTY, RESIDENTS, AND CLINICAL/RESEARCH FELLOWS MENTORED:
Graduate Students
PhD Students Advised
1984 Lee M. Greenberger, Photoreceptor Disc Shedding in Eye Cups, Emory School of Medicine
1987 Mary E. Pierce, Circadian Regulation of Cone Retinomotor Movement, Emory School of Medicine
1989 Cynthia Jo Horst, Microtubule-Membrane Assemblage of the Photoreceptor Connecting Cilium, Emory School of Medicine
1992 Michael S. Grace, Melatonin Degradation within the Vertebrate Eye and Pineal Gland, Emory School of Medicine
1993 Virgil Muresan, Molecular Dissection of the Photoreceptor Connecting Cilium Cytoskeleton: Biochemical Characterization of Basal Body and Axoneme-Plasmalemma Cross-Linkers in Relation to Structure and Function, University of Kansas School of Medicine
2000 Brooke Steenhard, Clock Genes and Clock-Controlled Genes in Retina: Studies on the Regulation of Period and Tryptophan Hydroxylase, University of Kansas School of Medicine
 
Postdoctoral Researchers
1981 - 1986 Dennis Defoe, Postdoctoral Fellow, Emory School of Medicine
1981 - 1982 Kamla Dutt, Research Associate, Emory School of Medicine
1983 - 1986 Brian Matsumoto, Research Associate, Emory School of Medicine
1986 - 1988 Andrew Sweatt, Postdoctoral Fellow, Emory School of Medicine
1987 - 1993 Gregory M. Cahill, Postdoctoral Fellow, Emory and University of Kansas School of Medicine
1990 - 1994 Menda Wetzel, Research Associate, University of Kansas School of Medicine
1991 - 1996 Carla Green, Postdoctoral Fellow, University of Kansas School of Medicine
1993 - 1995 Elena Bendala-Tufanisco, Postdoctoral Fellow, University of Kansas School of Medicine
1994 - 1996 David Osterbur, Research Associate, University of Kansas School of Medicine
1995 - 1997 Brian Hollander, Postdoctoral Fellow, University of Kansas School of Medicine
1996 - 2000 Mei-Ying Liang, Postdoctoral Fellow, University of Kansas School of Medicine and Medical College of Wisconsin
 

GRANT REVIEWS:
1979 NSF - Sensory Physiology. Ad hoc reviewer
1981 - 1986 *NIH Study Section: Regular Member, Vision Research Review Committee, National Eye Institute
1982 NIH/NEI Core Grant Site Visit: Illinois Eye and Ear Infirmary, University of Illinois
1983 NIH/NEI Core Grant Site Visit: Berman-Gunn Laboratory, Harvard Medical School
1984 NIH/NEI Core Grant Site Visit: Department of Ophthalmology, Yale University School of Medicine
1984 NIH/NEI Core Grant Site Visit: Institute of Biological Sciences, Oakland University, Rochester, MI
1985 NIH/NEI Core Grant Site Visit: Dept. of Anatomy, University of Pennsylvania
1988 NIH/NEI Core Grant Site Visit: Cullen Eye Institute, Baylor College of Medicine
1989 - 1991 *Study Section Member: Fight-for-Sight, Inc. Fellowship Review Panel
1990 Ad Hoc Member: Cell and Developmental Biology Study Section, American Cancer Society
1990 Ad Hoc Member; NIH Visual Sciences Study Section
1991 Chair (Ad Hoc): NIH Visual Sciences Study Section
1992 - 1996 *NIH Study Section: Regular Member, Visual Sciences C Study Section, Center for Scientific Review, National Institutes of Health
11/2002 Member: NIH Site visit team for the Laboratory of Developmental Neurobiology, National Institute of Child Health and Human Development
08/2005 Ad Hoc Member: NIH/NEI Review Group for Vision Science Training Grants
06/2007 Chair (Ad Hoc), NIH Study Section on Retinopathy Studies CB-G90. Center for Scientific Review
11/2007 Ad Hoc Member: NIH F05 panel for NRSA Fellowships in Cell Biology and Development, Center for Scientific Review
2008 Member, NEI Scientific Director Review Panel, National Institutes of Health
02/2009 Co-Chair (Ad Hoc), NIH Study Section on Retinopathy Studies CB-G90
06/2010 Chair (Ad Hoc), NIH Study Section on Retinopathy Studies CB-G90
04/2012 Ad Hoc Reviewer. Retinitis Pigmentosa Foundation of Canada
03/2014 NIH Study Section ZEY1 VSN 02. NEI Career Development and Pathways to Independence Applications
02/2015 Ad Hoc member, Biology of the Visual System (BVS) Study Section. Center for Scientific Review, National Institutes of Health
2016 Ad Hoc Member, NIH/NIDCR Study Section covering T32/T90/R90 Training Programs in Oral and Craniofacial Disease. National Institute of Dental Health and Craniofacial Disease.
2017 Ad Hoc Member. NEI T32/T35/K12 Review Committee, ZEY1 VSN (03). NEI Training Grants. October 2017.
2018 Ad Hoc Member. NEI K99 Review Committee, ZEY1VSN-K99. Research Career Transition Awards. March 2018


BIBLIOGRAPHY
Refereed Journal Publications/Original Papers
1. Besharse, J.C. 1971. Maturity and sexual dimorphism in the skull, mandible and teeth of the beaked whale, Mesoplodon densirostris. J. Mammal. 52: 297-315.
2. Besharse, J.C. and R.A. Brandon. 1973. Optomotor response and eye structure in the troglobitic salamander Gyrinophilus palleucus. Amer. Mid. Natur. 89: 463-467.
3. Besharse, J.C. and R.A. Brandon. 1974. Size and growth of the eyes of the troglobitic salamander Typhlotriton spelaeus. Inter. J. Speleol. 6: 255-264.
4. Besharse JC, Brandon RA. Postembryonic eye degeneration in the troglobitic salamander Typhlotriton spelaeus. J Morphol. 1974 Dec;144(4):381-405.
5. Besharse, J.C. and R.A. Brandon. 1976. Effects of continuous light and darkness on the eyes of the troglobitic salamander Typhlotriton spelaeus. J. Morphol. 149: 527-545.
6. Besharse, J.C. and Hollyfield, J.G. 1976. Renewal of normal and degenerating photoreceptor outer segments in the Ozark cave salamander. J. Exp. Zool. 198: 287-301.
7. Hollyfield, J.G., J.C. Besharse and M.E. Rayborn. 1976. The effect of light on the quantity of phagosomes in the pigment epithelium. Exp. Eye Res. 23: 623-635.
8. Bridges, C.D.B., J.G. Hollyfield, J.C. Besharse and M.E. Rayborn. 1976. Visual pigment loss after light-induced shedding of rod outer segments. Exp. Eye Res. 23: 637-641.
9. Besharse, J.C. and J.R. Holsinger. 1977. Gyrinophilus subterraneus: a new troglobitic salamander from West Virginia. Copeia 1977: 624-634.
10. Besharse, J.C. and J.G. Hollyfield. 1977. Ultrastructural changes during degeneration of photoreceptors and pigment epithelium in the Ozark Cave salamander. J. Ultrastructure Res. 59: 31-43.
11. Besharse, J.C. J.G. Hollyfield, and M.E. Rayborn. 1977. Photoreceptor outer segments: accelerated membrane renewal in rods after exposure to light. Science 196: 536-538.
12. Hollyfield, J.G., J.C. Besharse, and M.E. Rayborn. 1977. Variations in the phagosome content of the pigment epithelium in relationship to cyclic lighting. Atti. Fond. Giorgio Ronchi. 32: 309-313.
13. Hollyfield, J.G., J.C. Besharse, and M.E. Rayborn. 1977. Turnover of rod photoreceptor outer segments: membrane addition and loss in relationship to temperature. J. Cell Biol. 75: 409-506.
14. Besharse, J.C., J.G. Hollyfield and M.E. Rayborn. 1977. Turnover of rod photoreceptor outer segments: membrane addition and loss in relationship to light. J. Cell Biol. 75: 507-527.
15. Besharse, J.C. and J.G. Hollyfield. 1979. Turnover of mouse photoreceptor outer segments in constant light and darkness. Invest. Ophthalmol. Vis. Sci. 18: 1019-1024.
16. Danner, D.J., S.K. Lemmon, J.C. Besharse and L.J. Elsas, II. 1979. Purification and characterization of branched chain alpha-ketoacid dehydrogenase from bovine liver mitochondria. J. Biol. Chem. 254: 5522-5526.
17. Besharse, J.C. R.O. Terrill and D.A. Dunis. 1980. Light evoked disc shedding by rod photoreceptors in vitro: relationship to medium bicarbonate concentration. Invest. Ophthalmol. Vis. Sci. 19: 1512-1517.
18. Besharse, J.C. and K.H. Pfenninger. 1980. Membrane assembly in retinal photoreceptors. I. Freeze-fracture analysis of cytoplasmic vesicles in relationship to disc assembly. J. Cell Biol. 87: 451-463.
19. Besharse, J.C. D.A. Dunis and B. Burnside. 1982. Effects of adenosine 3', 5' cyclic monophosphate on photoreceptor disc shedding and retinomotor movement. Inhibition of rod shedding and stimulation of cone elongation. J. Gen. Physiol. 79: 775-790.
20. Besharse, J.C. 1982. The daily light-dark cycle and rhythmic metabolism in the photoreceptor-pigment epithelial complex. Progress Retinal Res. 1: 81-124.
21. Besharse, J.C. and D.A. Dunis. 1983. Rod photoreceptor disc shedding in eye cups: relationship to bicarbonate and amino acids. Exp. Eye Res. 36: 567-579.
22. Besharse, J.C. and D.A. Dunis. 1983. Methoxyindoles and photoreceptor metabolism: activation of rod shedding. Science 219: 1341-1343.
23. Iuvone, P.M. and J.C. Besharse. 1983. Regulation of indoleamine N-acetyltransferase activity in the retina: effects of light and dark, protein synthesis inhibitors and cyclic nucleotide analogs. Brain Research. 273: 111-119.
24. Besharse, J.C. and P.M. Iuvone. 1983. Circadian clock in the Xenopus eye controlling retinal serotonin N-acetyltransferase. Nature 305: 133-135.
25. Greenberger, L.M. and J.C. Besharse. 1983. Photoreceptor disc shedding in eye cups: inhibition by deletion of extracellular divalent cations. Investigative Ophthalmology Visual Science 24: 1456-1464.
26. Besharse, J.C., D.A. Dunis and P.M. Iuvone. 1984. Regulation and possible role of serotonin N-acetyltransferase in retina. Federation Proc. 43: 2704-2708.
27. Wood, J.G. and J.C. Besharse and L. Napier-Marshall. 1984. Partial characterization of lectin binding sites of retinal photoreceptor outer segments and interphotoreceptor matrix. Journal Comparative Neurology. 273: 299-307.
28. Defoe, D.M. and J.C. Besharse. 1985. Membrane assembly in retinal photoreceptors. II. Immunocytochemical analysis of freeze-fractured rod photoreceptor membranes using anti-opsin antibodies. Journal of Neuroscience. 5: 1023-1034.
29. Besharse, J.C., D.M. Forestner and D.M. Defoe. 1985. Membrane assembly in retinal photoreceptors. III. Distinct membrane domains of the connecting cilium of developing rods. Journal of Neuroscience. 5: 1035-1048.
30. Matsumoto, B. and J.C. Besharse. 1985. Light and temperature modulated staining of the rod outer segment distal tips with lucifer yellow. Invest. Ophthalmol. Vis. Sci. 26: 628-635.
31. Greenberger, L.M. and J.C. Besharse. 1985. Stimulation of photoreceptor disc shedding and pigment epithelial phagocytosis by glutamate, aspartate and other amino acids. J. Comp. Neurol. 239: 361-372.
32. Pierce, M.E. and J.C. Besharse. 1985. Circadian regulation of retinomotor movements: I. Interaction of melatonin and dopamine in the control of cone length. J. General Physiology. 86: 671-689.
33. Papermaster, D.S., B.G. Schneider and J.C. Besharse. 1985. Vesicular transport of newly synthesized opsin from the Golgi apparatus, toward the rod outer segment: Ultrastructural, immunocytochemical and autoradiographic evidence in Xenopus retinas. Invest. Ophthalmol. Visual Science. 26: 1386-1404.
34. Iuvone, P.M. and J.C. Besharse. 1986. Cyclic AMP stimulates serotonin N-acetyltransferase activity in Xenopus retina in vitro. J. Neurochem. 46: 33-39.
35. Iuvone, P.M. and J.C. Besharse. 1986. Involvement of calcium in regulation of serotonin N-acetyltransferase in retina. J. Neurochem. 46: 82-88.
36. Iuvone, P.M. and J.C. Besharse. 1986. Dopamine receptor-mediated inhibition of serotonin N-acetyltransferase activity in retina. Brain Res. 369: 168-176.
37. Papermaster, D.S., B.G. Schneider, D. Defoe and J.C. Besharse. 1986. Biosynthesis and vectorial transport of opsin on vesicles in retinal rod photoreceptors. J. Histochem. Cytochem. 34: 5-16.
38. Besharse, J.C., G. Spratt and D.M. Forestner. 1986. Light-evoked and kainic acid induced disc shedding by rod photoreceptors: differential sensitivity to extracellular calcium. J. Comp. Neurol. 251: 185-197.
39. Defoe, D.M., J.C. Besharse and S.J. Fliesler. 1986. Tunicamycin-induced dysgenesis of retinal rod outer segment membranes: II. Quantitative freeze-fracture analysis. Invest. Ophthalmol. Vis. Sci. 27: 1595-1601.
40. Matsumoto, B., D.M. Defoe and J.C. Besharse. 1987. Membrane turnover in rod photoreceptors: Ensheathment and phagocytosis of outer segment distal tips by pseudopodia of retinal pigment epithelium. Proc. Royal Soc. London B. 230: 339-354.
41. Pierce, M.E., and J.C. Besharse. 1987. Melatonin and rhythmic photoreceptor metabolism: melatonin-induced cone elongation is blocked at high light intensity. Brain Res. 405: 400-404.
42. Horst, C.J., D.M. Forestner, and J.C. Besharse. 1987. Cytoskeletal-membrane interactions: A stable interaction between cell surface glycoconjugates and doublet microtubules of the photoreceptor connecting cilium. J. Cell Biol. 105: 2973-2987.
43. Besharse, J.C. P.M. Iuvone, and M.E. Pierce. 1988. Regulation of rhythmic photoreceptor metabolism: A role for post-receptoral neurons. Progress Retinal Research 7: 21-61.
44. Sale, W.S., J.C. Besharse and G. Piperno. 1988. Distribution of acetylated alpha-tubulin in retina and in in vitro assembled microtubules. J. Cell Motility. 9: 243-253.
45. Pierce, M.E. and J.C. Besharse. 1988. Circadian regulation of retinomotor movements: II. The role of GABA in the Regulation of Cone Position. J. Comp. Neurol. 270: 279-287.
46. Besharse, J.C. and G. Spratt. 1988. Excitatory amino acids and rod photoreceptor disc shedding: Analysis Using Specific Agonists. Exp. Eye Res. 47: 609-620.
47. Besharse, J.C., G. Spratt and L. Reif-Lehrer. 1988. Effects of kynurenate and other excitatory amino acid antagonists as blockers of light- and kainate-induced retinal rod photoreceptor disc shedding. J. Comp. Neurol. 274: 295-303.
48. Witkovsky, P., S. Stone and J.C. Besharse. 1988. Dopamine modifies the balance of rod and cone inputs to horizontal cells of the Xenopus retina. Brain Res. 449: 332-336.
49. Witkovsky, P., S. Stone and J.C. Besharse. 1988. The effects of dopamine and related ligands on photoreceptor to horizontal cell signal transfer in the Xenopus retina. Biomedical Research. 9(Supplement): 93-107.
50. Cahill, G.M., and J.C. Besharse. 1989. Retinal melatonin is metabolized within the eye of Xenopus laevis. Proc. Nat'l. Acad. Sci. (USA). 86: 1098-1102.
51. Defoe, D.M., B. Matsumoto, and J.C. Besharse. 1989. Cytochalasin D inhibits glutamate-induced disc shedding without altering glutamate-induced increase in adhesivity. Exp. Eye Res. 48: 641-652.
52. Cahill, G.M. and J.C. Besharse. 1990. Circadian regulation of melatonin in the retina of Xenopus laevis: Limitation by Serotonin availability. J. Neurochem. 54:716-719.
53. Horst, C. J., L. V. Johnson and J. C. Besharse. 1990. Transmembrane assemblage of the photoreceptor connecting cilium and motile cilium transition zone contain a common immunologic epitope. Cell Motility and Cytoskeleton. 17: 329-344.
54. Cahill, G. M., M. S. Grace and J. C. Besharse. 1991. Rhythmic regulation of retinal melatonin: metabolic pathways, neurochemical mechanisms and the ocular circadian clock. Cellular and Molecular Pharmacology. 11: 529-560.
55. Cahill, G.M. and J.C. Besharse. 1991. Resetting the Circadian Clock in Cultured Xenopus Eyecups: Regulation of Retinal Melatonin Rhythms by Light and D2 Dopamine Receptors. J. Neuroscience. 11: 2959-2971.
56. Grace, M.S., G.M. Cahill and J.C. Besharse. 1991. Melatonin deacetylation: Retinal Vertebrate Class Distribution and Xenopus laevis Tissue Distribution. Brain Research. 559: 56-63.
57. Besharse, J.C. 1992. The "ON"-Bipolar Agonist, L-2-amino-4- phosphonobutyrate, Blocks Light-Evoked Cone Contraction in Xenopus Eyecups. Neurochemical Research. 17: 75-80.
58. Besharse, J.C. and P.M. Iuvone. 1992. Critique: Is Dopamine a Light-Adaptive or a Dark-Adaptive Modulator in Retina? Neurochemistry International. 20: 193-199.
59. Besharse, J.C. and P. Witkovsky. 1992. Light-Evoked Contraction of Red Absorbing Cones in the Xenopus retina is Maximally Sensitive to green Light. Visual Neuroscience. 8: 243-249.
60. Cahill, G. M. and J. C. Besharse. 1992. Light sensitive melatonin synthesis by Xenopus photoreceptors after destruction of the inner retina. Visual Neuroscience. 8: 487-490.
61. Defoe, D. M., B. Matsumoto and J. C. Besharse. 1992. Reconstitution of the Photoreceptor Pigment Epithelium Interface: L-Glutamate Stimulation of Adhesive Interactions and Rod Disc Shedding After Recombination of Dissociated Xenopus Eye Cups. Experimental Eye Research. 54: 903-911.
62. Grace, M. S. and J. C. Besharse. 1993. Solubilization and biochemical characterization of the melatonin deacetylase from Xenopus laevis retina. J. Neurochemistry. 60: 990-999.
63. Muresan, Z. and J. C. Besharse. 1993. D2-like dopamine receptors in amphibian retina: localization with fluorescent ligands. J. Comp. Neurology. 330: 1-12.
64. Muresan, V., H. C. Joshi, and J. C. Besharse. 1993 gamma-Tubulin in differentiated cell types: Localization in the vicinity of basal bodies in retinal photoreceptors and ciliated epithelia. J. Cell Science. 104: 1229-1237.
65. Wetzel, M. G., E. B. Tufanisco and J. C. Besharse. 1993. Tunicamycin does not inhibit transport of phosphatidylinositol to Xenopus rod outer segments. J. Neurocytology. 22: 397-412.
66. Cahill, G. M. and J. C. Besharse. 1993. Circadian clock functions localized in Xenopus retinal photoreceptors. Neuron. 10: 573-577.
67. Green, C. B and J. C. Besharse. 1994. Tryptophan hydroxylase expression is regulated by a circadian clock in Xenopus laevis retina. J. Neurochem. 62: 2420-2428.
68. Muresan, V. and J. C. Besharse. 1994. Complex intermolecular interactions maintain a stable linkage between the photoreceptor connecting cilium axoneme and plasma membrane. Cell Motility and Cytoskeleton. 28: 213-230.
69. Wetzel, M. G. and J. C. Besharse. 1994. Transport of phosphatidylcholine to Xenopus photoreceptor rod outer segments in the presence of tunicamycin. J. Neurocytology. 23: 333-342.
70. Grace, M. S. and J. C. Besharse. 1994. Melatonin deacetylase activity in the pineal gland and brain of the lizards Anolis carolinensis and Sceloporus jarrovi. Neuroscience. 62: 615-623.
71. Cahill, G. M. and J. C. Besharse. 1995. Circadian rhythmicity in vertebrate retinas: regulation by a photoreceptor oscillator. Progress Retinal Res. 14: 267-291.
72. Green, C. B., G. M. Cahill and J. C. Besharse. 1995. Tryptophan hydroxylase is expressed by photoreceptors in Xenopus leavis retina. Visual Neuroscience. 12: 663-670.
73. Green, C. B., G M. Cahill and J. C. Besharse. 1995. Regulation of tryptophan hydroxylase expression by a retinal circadian oscillator in vitro. Brain Research. 677: 283-290.
74. Besharse, J. C. and M. G. Wetzel. 1995. Immunocytochemical localization of opsin in rod photoreceptors during periods of rapid disc assembly. Journal of Neurocytology. 24: 371-388.
75. Green, C. B. and J. C. Besharse. 1996. Use of a high stringency differential display screen for circadian clock regulated retinal mRNAs. Molecular Brain Research. 37: 157-165.
76. Grace, M. S., L. A. Wang, G. E. Pickard, J. C. Besharse, and M. Menaker. 1996. The tau mutation shortens the period of rhythmic photoreceptor outer segment disk shedding in the hamster. Brain Research. 735: 93-100.
77. Green, C. B., J. C. Besharse and M. Zatz. 1996. Tryptophan hydroxylase mRNA levels are regulated by the circadian clock, temperature and cAMP in chick pineal cells. Brain Res. 738: 1-7.
78. Green, C. B. and J. C. Besharse. 1996. Identification of a novel vertebrate circadian clock regulated gene encoding the protein nocturnin. Proc. Nat. Acad. Sci. 93: 14884-14888.
79. Muresan, V., E. Bendala-Tufanisco, B. A. Hollander and J. C. Besharse. 1997. Evidence for kinesin-related proteins associated with the axoneme of retinal photoreceptors. Exp. Eye Research. 64: 895-903.
80. Green, C. B. and J. C. Besharse. 1997. Identification of vertebrate circadian clock-regulated genes by differential display. In: A. Pardee and P. Liang (eds). “Differential Display: Methods and Applications” in the series Methods in Molecular Biology. Vol 85: pp 219-230. The Humana Press, Inc., Totawa, N. J.
81. Knox, B. E., Schlueter, C., Steenhard, B. M., Green, C. B. and J. C. Besharse. 1998. Transgene expression in Xenopus rods. FEBS Letters. 423: 117-121.
82. Cahill, G. M., Parson, S. E. and J. C. Besharse. 1998. Spectral sensitivity of melatonin synthesis suppression in Xenopus eyecups. Visual Neuroscience. 15:499-502.
83. Hollander, B. A., Liang, M.-Y., and J. C. Besharse. 1999. Linkage of a nucleolin-related protein and casein kinase II with the detergent stable, photoreceptor cytoskeleton. Cell Motility and the Cytoskeleton. 43: 114-127.
84. Mani, S. S., Besharse, J. C. and B. E. Knox. 1999. Immediate upstream sequence of arrestin directs rod specific expression in Xenopus. Journal Biological Chemistry. 274: 15590-15597.
85. Green, C.B., Liang, M.-Y., Steenhard, B. M. And J. C. Besharse. 1999. Ontogeny of circadian and light regulation of melatonin release in Xenopus laevis embryos. Devel. Brain Res. 117: 109-116
86. Zhuang, M., Y. Wang, B. M. Steenhard, and J. C. Besharse. 2000. Differential regulation of two period genes in the Xenopus eye. Molecular Brain Research. 82: 52-64.
87. Steenhard, B. M. and J. C. Besharse. 2000. Phase shifting the retinal circadian clock: xPer2 mRNA induction by light and dopamine. Journal of Neuroscience. 20: 8572-8577.
88. Besharse, J. C. 2001. Preview: Coupling an activated MAP kinase pathway to circadian output clock. Neuron. 29: 3-4.
89. Wang, Y, Osterbur, D. L., Megaw, P. L., Tosini, G., Fukuhara, C., Green, C. B., and J. C. Besharse. 2001. Rhythmic Expression of Nocturnin mRNA in Multiple Tissues of the Mouse. BMC Developmental Biology. 1: 9.
90. Kita, Y., M. Shiozawa, W. Jin, R. R. Majewsky, J. C. Besharse, A. S. Greene, and H. J. Jacob. 2002. Implications of circadian gene expression in in kidney and liver and the effects of fasting on pharmacogenetic studies. Parmacogenetics. 12: 55-65.
91. Pazour, G.L., S.A. Baker, J.A. Deane, D.G., Cole, B.A. Dickert, J.L. Rosenbaum, G.B. Witman, and J.C. Besharse. 2002. The intraflagellar transport protein, IFT88, is essential for photoreceptor assembly and maintenance. J. Cell Biology. 157: 103-114.
92. Mikami, A., S.H. Tynan, T. Hama, K. Luby-Phelps, T. Saito, J.E. Crandall, J.C. Besharse, R.B. Vallee. 2002. Molecular structure of cytoplasmic dynein 2 and its distribution in neuronal and ciliated cells. J. Cell Science. 120: 4801-4808.
93. Luby-Phelps, K., G. Ning, J. Fogerty and J.C. Besharse. 2003. Visualization of identified GFP-expressing cells at the light and electron microscopic level. J. Histochem. and Cytochem. 51: 271-274.
94. Besharse, J.C., S.A. Baker, K. Luby-Phelps and G. J. Pazour. 2003. Photoreceptor intersegmental transport and retinal degeneration, a conserved pathway common to motile and sensory cilia. Adv Exp Med Biol. 2003: 533:157-64.
95. Baker, S.A., K. Freeman, K. Luby-Phelps, G.J. Pazour, and J.C. Besharse. 2003. IFT20 Links Kinesin II with a Mammalian Intraflagellar Transport Complex that is Conserved in Motile Flagella and Sensory Cilia. J. Biological Chemistry. 278: 34211-34218.
96. Green C.B. and J.C. Besharse. 2004. Retinal Circadian Clocks and Control of Retinal Physiology. J. Biological Rhythms 9: 91-102.
97. Peet, J., Bragin, A., Calvert, P, S., Nikonov, S.S., Mani, S., Zhao, J., Besharse, J.C., Pierce, E., Knox, B., and E. N. Pugh, Jr. 2004. Cytoplasmic Spaces of Living CHO and Rod Photoreceptor Cells Visualized and Quantified with EGFP. J. Cell Science. 117: 3049-3059.
98. Besharse, J.C., Zhuang, M., Freeman, K. and J. Fogerty. 2004. Regulation of Photoreceptor Per1 and Per2 by Light, Dopamine, and a Circadian Clock. European Journal of Neuroscience. 20: 167-174.
99. Gabarino-Pico, E., S. Niu, M. D. Rollag, C. A. Strayer, J.C. Besharse, and C.B. Green. 2007. Immediate early responses of the circadian poly A ribonuclease nocturnin to two extracellular stimuli. RNA. 13: 745-755. PMCID: PMC1852813.
100. Green, C.B., Douris, N., Kojima, S, Strayer, C.A., Fogerty, J., Lourim, D., Keller, S.A., and J. C. Besharse. 2007. Loss of nocturnin, a circadian deadenylase, confers resistance to hepatic steatosis and diet-induced obesity. Proceedings National Academy Sciences. 104: 9888-9893. PMCID: PMC1871564
101. Luby-Phelps, K., Fogerty, J., Baker, S. A., Pazour, G. J., and J. C. Besharse. 2008. Spatial Distribution of Intraflagellar Transport Proteins in Vertebrate Photoreceptors. Vision Research. 48: 413-423. PMCID: PMC2276311
102. Insinna, C., Pathak, N., Perkins, B., Drummond, I., and J. C. Besharse. 2008. The Homodimeric Kinesin, Kif17, Is essential for Photoreceptor Outer Segment Development. Developmental Biology. 316: 160-170. PMCID: PMC2362383
103. Insinna, C. and J. C. Besharse. 2008. Intraflagellar Transport and the Sensory Outer Segment of Vertebrate Photoreceptors. Developmental Dynamics. 237: 1982-1992. PMCID: PMC2692564
104. Bhowmick, R., Li, M., Sun, J., Baker, S. A., Insinna, C. and J. C. Besharse. 2009. Photoreceptor IFT Complexes Containing Chaperones, Guanylyl Cyclase 1, and Rhodopsin. Traffic. 10: 648-663. PMCID: PMC2827254
105. Insinna, C., Humby, M., Sedmak, T., Wolfrum, U., and J. C. Besharse. 2009. Different Roles for KIF17 and Kinesin II in Photoreceptor Development and Maintenance. Developmental Dynamics. 238: 2211-2222. PMCID: PMC2733920.
106. Insinna, C., Luby-Phelps, K., Link, B. A. and J. C. Besharse. 2009. Analysis of IFT Kinesins in Developing Zebrafish Cone Photoreceptor Sensory Cilia. Methods in Cell Biology. 93: 219- 234.
107. Insinna, C., Baye, L.M., Amsterdam, A., Besharse, J.C. and B.A. Link. 2010. Analysis of a zebrafish dync1h1 mutant reveals multiple functions for cytoplasmic dynein 1 during retinal photoreceptor development. Neural Development. 5: 12.
108. Boye, S.E., Boye, S.L., Pang, J., Ryals, R., Everhart, D., Umino, Y., Neeley, A.W., Besharse, J.C., Barlow, R., and W.W. Hauswirth. 2010. Functional and behavioral restoration of vision by gene therapy in a mouse model of Leber congenital amaurosis-1 (LCA1). PLoS One 5(6): e11306.
109. Fogerty, J. and J.C. Besharse. 2011. 174delG mutation in mouse MFRP causes photoreceptor degeneration and RPE atrophy. Investigative Ophthalmology and Visual Science. 50: 7256-7266. (published online August 2, 2011). PMID: 21810984.
110. Scoma, H.D., Humby, M., Yadav, G. Zhang, Q., Fogerty, J. and J.C. Besharse. 2011. The de-ubiquitinylating enzyme, USP2, is associated with the circadian clockwork and regulates its sensitivity to light. PLoS One. 6(9): e25382. PMID: 21966515.
111. Wong-Riley, M.T.T. and J.C. Besharse. 2012. The kinesin superfamily protein KIF17: one protein with many functions. Biomolecular Concepts. 3(3): 267–282.
112. Malicki, J. and J.C. Besharse. 2012. Kinesin-2 family motors in the unusual photoreceptor cilium. Vision Research. 75: 33-36. PMID: 23123805.
113. Bader, J.R., Kusik, B.W. and J.C. Besharse. 2012. Analysis of KIF17 distal tip trafficking in zebrafish cone photoreceptors. Vision Research. 75: 37-43. PMID: 23099049.
114. Fogerty, J. and J.C. Besharse. 2014. Subretinal infiltration of monocyte derived cells and complement misregulation in mice with AMD-like pathology. Advances Experimental Medicine and Biology. 801: 355-363.
115. Miesfeld JB, Gestri G, Clark BS, Flinn MA, Poole RJ, Bader JR, Besharse JC, Wilson SW, Link BA. Yap and Taz regulate retinal pigment epithelial cell fate. Development. 2015 Sep 01;142(17):3021-32. PMCID: PMC4582179
116. Besharse JC, McMahon DG. The Retina and Other Light-sensitive Ocular Clocks. J Biol Rhythms. 2016 Jun;31(3):223-43. PMCID: PMC5479307
117. Collery RF, Volberding PJ, Bostrom JR, Link BA, Besharse JC. Loss of Zebrafish Mfrp Causes Nanophthalmia, Hyperopia, and Accumulation of Subretinal Macrophages. Invest Ophthalmol Vis Sci. 2016 Dec 01;57(15):6805-6814. PMCID: PMC5215506
118. Lewis TR, Kundinger SR, Pavlovich AL, Bostrom JR, Link BA, Besharse JC. Cos2/Kif7 and Osm-3/Kif17 regulate onset of outer segment development in zebrafish photoreceptors through distinct mechanisms. Dev Biol. 2017 May 15;425(2):176-190. PMCID: PMC5558849
119. Lewis TR, Zareba M, Link BA, Besharse JC. Cone myoid elongation involves unidirectional microtubule movement mediated by dynein-1. Mol Biol Cell. 2018 Jan 15;29(2):180-190. PMCID: PMC5909930
120. Lewis TR, Kundinger SR, Link BA, Insinna C, Besharse JC. Kif17 phosphorylation regulates photoreceptor outer segment turnover. BMC Cell Biol. 2018 Nov 20;19(1):25. PMCID: PMC6245759
121. Han DP, Skumatz C, Besharse JC, Kassem IS. Sustained Anti-Vascular Endothelial Growth Factor Activity of Aflibercept (Eylea) After Storage in Polycarbonate Syringes Used for Intravitreal Injection: A Pathway to Safety and Efficiency. J Ocul Pharmacol Ther. 2019 Jun;35(5):278-282. PMCID: PMC6588124
122. Sadler KE, Lewis TR, Waltz TB, Besharse JC, Stucky CL. Peripheral nerve pathology in sickle cell disease mice. Pain Rep. 2019 Jul-Aug;4(4):e765. PMCID: PMC6728004
123. Spencer WJ, Ding JD, Lewis TR, Yu C, Phan S, Pearring JN, Kim KY, Thor A, Mathew R, Kalnitsky J, Hao Y, Travis AM, Biswas SK, Lo WK, Besharse JC, Ellisman MH, Saban DR, Burns ME, Arshavsky VY. PRCD is essential for high-fidelity photoreceptor disc formation. Proc Natl Acad Sci U S A. 2019 Jun 25;116(26):13087-13096. PMCID: PMC6601265
 
Books, Chapters, and Reviews
1. Besharse, J.C. 1980. Light and membrane biogenesis in rod photoreceptors of vertebrates. In: The Effects of Constant Light on Visual Processes. T. Williams and B. Baker, eds., Plenum Press, New York, pp. 409-431.
2. Besharse, J.C. and D.A. Dunis. 1982. Rod photoreceptor disc shedding in vitro: Inhibition by Cytochalasins and Activation by Colchicine. In: The Structure of the Eye IV, J.G. Hollyfield, ed., Elsevier-North Holland, New York, pp. 85-96.
3. Besharse, J.C. 1986. Photosensitive membrane turnover: differentiated membrane domains and cell-cell interaction. In "The Retina A Model for Cell Biological Studies Part I." (R. Adler and D. Farber, eds.). pp. 297-352. Academic Press, Inc. New York.
4. Pierce, M.E. and J.C. Besharse. 1986. Melatonin and dopamine interactions in the regulation of rhythmic photoreceptor metabolism. In: "Pineal-Retina Relationships" pp.219-237. (P. O'Brien and D. Klein, eds.). Academic Press, New York.
5. Besharse, J.C. and C.J. Horst. 1990. The photoreceptor connecting cilium: A model for the transition zone. In "Structure and Function of Ciliary and Flagellar Surfaces." (R.A. Bloodgood, ed.). Plenum Press, New York. pp. 389-417.
6. Besharse, J. C. and C. B. Green. 1995. The photoreceptor circadian clock: from disc shedding to gene regulation. pp. 9-23. In: Y. Christen, M. Doly, and M. T. Droy-Lefaix, eds. Les Séminaires Ophthalmologiques d’Ipsen, Tome 7. “Développment et Différentiation of the Rétine.” Irvinn, Paris.
7. Besharse, J. C. and D. M. Defoe. 1998. Role of the pigment epithelium in photoreceptor membrane turnover. In: M. F. Marmor and T. J. Wolfensberger, eds. “The Retinal Pigment Epithelium” (2nd Edition). Oxford University Press. (pp. 152-172).
8. Green, C. B. and J. C. Besharse. 1998. Molecular Analyses of the Xenopus Photoreceptor Circadian Oscillator. In: Biological Clocks. Mechanisms and Applications, Y. Touitou, ed. Elsevier Science B.V., Amsterdam.
9. Besharse, J. C. and C. B. Green. 1998. Gene regulation and the photoreceptor circadian clock. pp 33-45 in “Circadian Clocks and Entrainment” (K. Honma and S. Honma, eds). Hokkaidao Univ. Press, Sapporo, Japan.
10. Baker, S.A., Pazour, G.J., Witman, G.B., and J.C. Besharse. 2004. Photoreceptors and Intraflagellar Transport. Pp. 109-132 In: D.A. Williams (editor). Photoreceptor Cell Biology and Inherited Retinal Degenerations. Recent Advances in Human Biology, Vol. 10. WORLD SCIENTIFIC PUBLISHING COMPANY PTE LTD, Singapore.
11. Besharse, J. C. and C. Insinna. 2010. The Photoreceptor Outer Segment as a Sensory Cilium. In: Dartt, D.A., Besharse, J.C. and Dana, R. Eds. Encyclopedia of the Eye. Vol 3. Oxford: Academic Press, pp. 240-346.
12. Dartt, D.A., Besharse, J.C. and Dana, R. (Eds). 2010. Encyclopedia of the Eye. Volumes 1-4. Elsevier: Academic Press.
13. Besharse, J.C. and D. Bok (Eds). 2011. The Retina and its Disorders. Elsevier: Academic Press. 912 pages.
 
Non-Refereed Journal Publications/Original Papers
1. Lewis, T.R. Kundinger, S.R., Link, B.A., Insinna, C. and J.C. Besharse. 2018. Kif17 phosphorylation regulates its ciliary localization and photoreceptor outer segment turnover. bioRxiv 351122; doi: https://doi.org/10.1101/351122
2. Boye SE, Boye SL, Pang J, Ryals R, Everhart D, Umino Y, Neeley AW, Besharse J, Barlow R, Hauswirth WW. Functional and behavioral restoration of vision by gene therapy in the guanylate cyclase-1 (GC1) knockout mouse. PLoS One. 2010 Jun 25;5(6):e11306. PMCID: PMC2892468
3. Insinna C, Baye LM, Amsterdam A, Besharse JC, Link BA. Erratu to Analysis of a zebrafish dync1h1 mutant reveals multiple functions for cytoplasmic dynein 1 during retinal photoreceptor development [Neural Development, (2010), 5, 12] Neural Development. 6 August 2010;5(1).
4. Hollandei BA, Besharse JC. CO-purification of a casein k1nase I-like kinase and two prominant substrate phosphoproteins with the ciliary axoneme of bovine photoreceptor outer segments Investigative Ophthalmology and Visual Science. 1997;38(4).
5. Besharse JC. The photoreceptor circadian clock and regulation of gene expression Investigative Ophthalmology and Visual Science. 15 February 1996;37(3).
6. Green CB, Besharse JC. Identification of nocturnin, a novel photoreceptor specific leucine zipper protein that is regulated by a retinal circadian clock Investigative Ophthalmology and Visual Science. 15 February 1996;37(3).
7. Sanger BM, Green CB, Besharse JC. Characterization of the Xenopus tryptophan hydroxylase gene Investigative Ophthalmology and Visual Science. 15 February 1996;37(3).
8. Besharse JC, Parsons SE, Green CB. Protein synthesis inhibitors cause phase-dependent phase shifts of the photoreceptor circadian clock Investigative Ophthalmology and Visual Science. 15 February 1996;37(3).
9. Cahill GM, Besharse JC. Circadian rhythmicity in vertebrate retinas: Regulation by a photoreceptor oscillator Progress in Retinal and Eye Research. 1995;14(1):267-291.
10. Witkovsky P, Stone S, Besharse JC. The effects of dopamine and related ligands on photoreceptor to horizontal cell transfer in the Xenopus retina Biomedical Research. 1988;9(SUPPL. 2):93-107.
11. Besharse JC, Iuvone PM, Pierce ME. Chapter 2 Regulation of rhythmic photoreceptor metabolism: A role for post-receptoral neurons Progress in Retinal Research. 1988;7(C):21-61.
12. Besharse JC, Dunis DA. Methoxyindoles and photoreceptor metabolism: Activation of rod shedding Science. 1983;219(4590):1337-1339.
13. Besharse JC. Chapter 3 The daily light-dark cycle and rhythmic metabolism in the photoreceptor-Pigment epithelial complex Progress in Retinal Research. 1982;1(C):81-124.
14. Besharse JC. Photoreceptor shedding and retinomotor movement in vitro: Modulation by cyclic nucleotides Journal of Cell Biology. 1981;91(2 II).
15. Besharse C, Terrill RO, Dunis DA. Light-evoked disc shedding by rod photoreceptors in vitro: relationship to medium bicarbonate concentration. Invest Ophthalmol Vis Sci. 1980 Dec;19(12):1512-7.
16. Besharse JC, Terrill RO. Light-evoked shedding by rods in vitro: Activation by colchicine Investigative Ophthalmology and Visual Science. 1980;19(Suppl. 4):161.
17. Hollyfield JG, Besharse JC, Rayborn ME. Variations in the phagosome content of the pigment epithelium in relationship to cyclic lighting Atti della Fondazione Giorgio Ronchi. 1977;32(2):309-313.
18. Besharse JC, Insinna C. The photoreceptor outer segment as a sensory cilium Encyclopedia of the Eye, Four-Volume Set. 1 January 2010:340-346.
19. Besharse JC, Insinna C. The photoreceptor outer segment as a sensory cilium Encyclopedia of the Eye. 1 January 2010:340-346.