Jonathan M. Scholey

image of Jonathan M. Scholey

Distinguished Professor Emeritus of Cell Biology and Biochemistry


Molecular & Cellular Biology

Offices and Labs

3253 Life Sciences Addition

Profile Introduction

My research has focused on the cell biology of motor proteins i.e. subcellular processes that depend upon the action of cytoskeletal motors. I studied myosin motors as a PhD student with Dr Jake Kendrick-Jones at the MRC Laboratory of Molecular Biology, Cambridge and mitotic motors (kinesins and dyneins) as a post-doctoral fellow with Dr Dick McIntosh at CU Boulder. This provided the foundation for my lab's research on; (i) Intraflagellar transport motors in ciliogenesis i.e. heterotrimeric/homodimeric kinesin-2 motors and IFT-dynein; and (ii) Mitotic motors and chromosome segregation e.g. bipolar kinesin-5 motors and the sliding filament mechanism of mitotic spindle elongation (anaphase B). Regarding teaching and education, I have a long-standing interest in science at the interface of cell biology and physical sciences. Accordingly, after obtaining a BSc in cell and molecular biology from the Biophysics Department at Kings College, London University and a PhD in molecular biology from the MRC LMB/Trinity College, Cambridge University, as a professor at UC Davis, I taught various classes in biochemistry, biophysics, cell and molecular biology, including an undergraduate cell biology class based on the iBioseminars series which I worked on with members of the iBiology team. I also taught similar classes as a Fulbright Scholar and Adjunct Professor in the Department of Molecular Biology and Genetics at Bosphorus University in Istanbul, Turkey


1981 PhD Molecular Biology MRC Laboratory of Molecular Biology and Trinity College, Cambridge University:
1977 BSc Cell and Molecular Biology (1st class honours) MRC Cell Biophysics Unit, King's College Department of Biophysics, London University:

Research Interests

Cell Biology: Motor Proteins in Mitosis and Ciliogenesis.

Motor Protein Functions in Mitosis and Intraflagellar Transport. Specific Projects: 1. Mechanisms of mitosis and chromosome segregation in the Drosophila embryo (funded by NIH grant GM55507): 2. Kinesin-2 motors, intraflagellar transport and sensory cilium biogenesis in C. elegans neurons (funded by NIH grant GM50718). Technical approaches; motor protein/nucleotide biochemistry and molecular biology; quantitative modelling (led by Dr G Civelekoglu-Scholey); light microscopy and the elucidation of motor/polymer dynamics and function in living cells.


1973 Chemistry GCE S-level (scholarship level), Northampton Grammar School.
1976 Sambrooke Exhibition in Natural Sciences, Department of Biophysics, Kings College, London.
1977-1980 MRC Studentship for training in molecular biology research methods under Dr J Kendrick-Jones in Dr HE Huxley's section of the MRC LMB (PhD thesis, 1981: Regulation of vertebrate nonmuscle and smooth muscle myosin filament assembly and interaction with actin by light chain phosphorylation).
1982-1985 MRC Travelling Postdoctoral Fellowship and BHF/AHA British-American Exchange Postdoctoral Fellowship for cell biology research with Dr JR McIntosh in MCDB at CU Boulder (Post-doctoral research: Motor Proteins in Mitosis).
1986-1988 Basil O'Connor Starter Scholar Research Award, March of Dimes Birth Defects Foundation.
1993 Lakian Summer Research Fellowship, Marine Biological Laboratory, Woods Hole, Mass.
1995 Honorary Member of the Golden Key National Honor Society
1996 Elected Chairperson, Gordon Research Conference on Motile and Contractile Systems.
2010-2011 Fulbright Scholar: Senior Lecturer Award in Molecular biology at Bogazici University, Istanbul, Turkey.
2014 Faculty Lifetime Achievement Award from the Biochemistry, Molecular, Cellular and Developmental Biology (BMCDB) Graduate Group at UC Davis.
2014-present Distinguished Professor (Emeritus 2015), Dept of Molecular and Cell Biology, UC Davis.
2017 Elected Fellow of the American Society for Cell Biology.

Department and Center Affiliations

1993-2015: Professor, Department of Molecular and Cell Biology, and Cell Biologist and Biochemist, Agriculture Experiment Station,UC Davis. Primary Appointment, Center for Genetics and Development.
2011-2017: Adjunct Teaching Professor, Department of Molecular Biology and Genetics, Bogazici University, Istanbul, Turkey.
2010 -2011: Visiting Professor and Fulbright Senior Lecturer in Molecular Biology, Department of Molecular Biology and Genetics, Bogazici University, Istanbul, Turkey.
2001-2006: Professor, Center for Genetics and Development, UC Davis.
1997-1998: Visiting Professor, Department of Biochemistry and Biophysics, UC San Francisco.
1989-1993: Assistant Professor, Department of Zoology, and Cell Biologist and Biochemist, Agriculture Experiment Station, UC Davis.
1986-1993: Assistant Professor, Departments of (i) Biochemistry, Biophysics and Genetics, and (ii) Cell and Structural Biology, University of Colorado Health Sciences Center, Denver, Colorado.
1986-1989: Staff Scientist, Department of Molecular and Cell Biology, National Jewish Hospital and Research Center, Denver, Colorado.
1982-1986: Post-doctoral Fellow, Department of Molecular, Cellular and Developmental Biology, University of Colorado, Boulder, Colorado.


American Society for Cell Biology, Emeritus Member.
American Society for Biochemistry and Molecular Biology (Member 2000-2007)
Faculty of 1000: Cell Biology faculty (Member 2005-2008)
Editorial Board Member, Journal of Biological Chemistry (2000-2007)
Editorial Board Member, Cytoskeleton
Editorial Board Member, Molecular Biology of the Cell
Biochemical Society (UK) Member
Member, NIH Cell Biology Integrated Review Group, Nuclear and Cytoplasmic Structure/Function and Dynamics Study Section (Member 2012 -2015)


Scholey Lab website

Teaching Interests

Biochemistry, biophysics, cell and molecular biology. Physics and chemistry of the cell.


MCB 110V iBioseminars in Cell and Molecular Biology ((Spring Quarter))
BIO 494 Quantitative Cell and Molecular Biology ((Fall 2013: Bogazici University Molecular Biology and Genetics Major))
MCB 143 Cell and Molecular Biophysics ((2001-2012))
MCB 212 Cell Biology ((Winter 2012))
MCB 221D Cellular Biochemistry ((1991-2012))
BIO 580 Molecular Biology of Mitosis and Cell Division ((Fall 2010: Bogazici University Molecular Biology and Genetics Graduate Group))
BIO 495 Cell and Molecular Biophysics ((Fall 2010: Bogazici University Molecular Biology and Genetics Major))
BIS 102 Biochemistry: Structure and Function of Biomolecules ((1993-2000))
BIS 104 Cell Biology ((1998-2000))
MCB 142 Advanced Cell Biology ((1990-1993))
BIS 1A Introductory Molecular Biology ((1989-1992))


8/6/2017 4:59:09 AM
  • Scholey JM, Taylor KA, Kendrick-Jones J. 1980. Regulation of non-muscle myosin assembly by calmodulin-dependent light chain kinase. Nature, 287: 233-235.

  • Kendrick-Jones J and Scholey JM. 1981. Myosin-linked Regulatory Systems. Journal of Muscle Research and Cell Motility, 2: 347-372.

  • Kendrick-Jones J, Cande WZ, Tooth PJ, Smith RC, Scholey JM. 1983. Studies on the effect of phosphorylation of the 20,000 Mr light chain of vertebrate smooth muscle myosin. Journal of Molecular Biology;165:139-62.

  • Scholey JM, Neighbors B, McIntosh JR, Salmon ED.1984. Isolation of microtubules and a dynein-like MgATPase from unfertilized sea urchin eggs. Journal of Biological Chemistry, 259: 6516-6525.

  • Scholey JM, Porter ME, Grissom PM, McIntosh, JR. 1985. Identification of kinesin in sea urchin eggs and evidence for its localization in the mitotic spindle. Nature, 318: 483-485.
  • Scholey JM, Heuser J, Yang JT, Goldstein LS. 1989. Identification of globular mechanochemical heads of kinesin. Nature, 338: 355-357.

  • Sawin KE and Scholey JM. 1991. Motor proteins in cell division. Trends in Cell Biology, 1:122-129.

  • Wright BD, Terasaki M, Scholey JM. 1993. Roles of kinesin and kinesin-like proteins in sea urchin embryonic cell division: evaluation using antibody microinjection. Journal of Cell Biology, 123: 681-689.

  • Cole DG, Chinn SW, Wedaman KP, Hall K, Vuong T, Scholey JM. 1993. Novel heterotrimeric kinesin-related protein purified from sea urchin eggs. Nature, 366: 268-270.
  • Cole DG, Saxton WM, Sheehan KB, Scholey JM. 1994. A "slow" homotetrameric kinesin-related motor protein purified from Drosophila embryos. Journal of Biological Chemistry. 269: 22913-22916.

  • Kashina AS, Baskin RJ, Cole DG, Wedaman KP, Saxton WM, Scholey JM. 1996. A bipolar kinesin. Nature, 379:270-272.

  • Wedaman KP, Meyer DW, Rashid DJ, Cole DG, Scholey JM. 1996. Sequence and submolecular localization of the 115-kD accessory subunit of the heterotrimeric kinesin-II (KRP85/95) complex. Journal of Cell Biology, 132: 371-380.

  • Scholey JM. 1996. Kinesin-II, a membrane traffic motor in axons, axonemes, and spindles. Journal of Cell Biology, 133:1-4.

  • Morris RL and Scholey JM. 1997. Heterotrimeric Kinesin-II Is Required for the Assembly of Motile 9+2 Ciliary Axonemes on Sea Urchin Embryos. Journal of Cell Biology, 138: 1009-1022.
  • Orozco JT, Wedaman KP, Signor D, Brown H, Rose L, Scholey JM. 1999. Movement of motor and cargo along cilia. Nature. 398: 674.

  • Signor D, Wedaman KP, Orozco JT, Dwyer ND, Bargmann CI, Rose LS, Scholey JM. 1999. Role of a class DHC1b dynein in retrograde transport of IFT motors and IFT raft particles along cilia, but not dendrites, in chemosensory neurons of living Caenorhabditis elegans. Journal of Cell Biology, 147: 519-530. 

  • Sharp DJ, Yu KR, Sisson JC, Sullivan W, Scholey JM. 1999. Antagonistic microtubule-sliding motors position mitotic centrosomes in Drosophila early embryos. Nature Cell Biology. 1: 51-54

  • Sharp DJ, Rogers GC, Scholey JM. 2000. Cytoplasmic dynein is required for poleward chromosome movement during mitosis in Drosophila embryos. Nature Cell Biology; 2:922-30.

  • Scholey JM. 2003. Intraflagellar transport. Annual Review of Cell and Developmental Biology. 19: 423-443.

  • Rogers GC, Rogers SL, Schwimmer TA, Ems-McClung SC, Walczak CE, Vale RD, Scholey JM, Sharp DJ. 2004. Two mitotic kinesins cooperate to drive sister chromatid separation during anaphase. Nature. 427: 364-370.

  • Brust-Mascher. I, Civelekoglu-Scholey G, Kwon M, Mogilner A,  Scholey JM, 2004, Model for Anaphase B: role of three mitotic motors in a switch from poleward flux to spindle elongation. Proc. Natl. Acad Sci USA. 101: 15938-15943.

  • Snow JJ, Ou G, Gunnarson AL, Walker MR, Zhou HM, Brust-Mascher I, Scholey JM. 2004. Two anterograde intraflagellar transport motors cooperate to build sensory cilia on C. elegans neurons. Nature Cell Biology. 6:1109-13.

  • Ou G, Blacque OE, Snow JJ, Leroux MR, Scholey JM. 2005. Functional coordination of intraflagellar transport motors. Nature. 436: 583-587.

  • Pan, X, Ou, G, Civelekoglu-Scholey, G, Blacque, O, Endres, N, Tao, L, Mogilner, A, Leroux, M, Vale, RD and Scholey, JM. 2006. Mechanism of Transport of IFT particles in C. elegans cilia by the concerted action of kinesin-II and OSM-3 motors. Journal of Cell Biology, 174: 1035-1045.
  • Civelekoglu-Scholey G, Sharp DJ, Mogilner A, Scholey JM. 2006. Model of chromosome motility in Drosophila embryos: adaptation of a general mechanism for rapid mitosis. Biophysical Journal, 90: 3966-82.

  • Goshima G, Scholey JM. 2010. Control of mitotic spindle length. Annual Review of Cell and Developmental Biology, 26: 21-57.

  • Hao L, Thein M, Brust-Mascher I, Civelekoglu-Scholey G, Lu Y, Acar S, Prevo B, Shaham S, Scholey JM. 2011. Intraflagellar transport delivers tubulin isotypes to sensory cilium middle and distal segments. Nature Cell Biology. 13:790-798.

  • Acar, S., Carlson, D.B, Budamagunta, M.S., Yarov-Yarovoy, V., Correia, J.J., Ninonuevo, M.R., Jia, W., Tao, L., Leary, J.A., Voss, J.C., Evans, J.E. and J.M. Scholey. 2013. The bipolar assembly domain of the mitotic motor kinesin-5. Nature Communications, 4: 1343.

  • Wang H, Brust-Mascher I, Civelekoglu-Scholey G, Scholey JM. 2013. Patronin mediates a switch from kinesin-13-dependent poleward flux to anaphase B spindle elongation. Journal of Cell Biology, 203: 35-46

  • Scholey, JM. 2013. Kinesin-2: A Family of Heterotrimeric and Homodimeric Motors with Diverse Intracellular Transport Functions. Annual Review of Cell and Developmental Biology. 29:443-469.

  • Prevo, B., Mangeol, P., Oswald, F., Scholey, JM and EJG Peterman. 2015. Functional differentiation of cooperating kinesin-2 motors orchestrates cargo import and transport in C. elegans cilia. Nature Cell Biology, 17: 1536-1545.

  • Brust-Mascher, I, Civelekoglu-Scholey, G. and JM Scholey. 2015. Mechanism for Anaphase B: Evaluation of "Slide-and-Cluster" versus "Slide-and-Flux-or-Elongate" Models. Biophysical Journal, 108: 2007-2018.

  • Scholey JM, Civelekoglu-Scholey G., and I Brust-Mascher. 2016. Anaphase B. Biology (Basel), 5(4). pii: E51.