Jonathan M. Scholey

Jonathan Scholey

Position Title
Distinguished Professor Emeritus

Department of Molecular and Cellular Biology, College of Biological Sciences

3253 Life Sciences Addition

Profile Introduction

My research has focused on the cell biology of motor proteins i.e. subcellular processes that depend on cytoskeletal motors. I studied myosin motors as a PhD student with Dr Jake Kendrick-Jones at the MRC LMB and mitotic motors (kinesins and dyneins) as a post-doc with Dr Dick McIntosh in MCDB at CU Boulder. This provided the foundation for my lab's research on; (i) Intraflagellar transport motors in ciliogenesis; and (ii) Mitotic motors in spindle assembly and anaphase B chromosome segregation. 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 Dept at KCL and a PhD in molecular biology from the MRC LMB/Trinity College, Cambridge, I taught various classes in biochemistry, biophysics, cell and molecular biology at UC Davis and at Bosphorus University in Istanbul.

Research Interests

Cell Biology: Motor Proteins in Mitosis and Ciliogenesis.

Specific Projects: 1. Mechanisms of mitosis and chromosome segregation in Drosophila and sea urchin embryos, notably bipolar kinesin-5 motors and the sliding filament mechanism of mitotic spindle elongation during anaphase (funded by NIH grant GM55507): 2. Intraflagellar transport (IFT) motors, notably heterotrimeric/homodimeric kinesin-2 and IFT-dynein, in IFT and cilium biogenesis in C. elegans sensory neurons and sea urchin swimming blastulae (funded by NIH grant GM50718). Technical approaches; protein/nucleotide biochemistry and molecular biology; motor protein purification and motility/gliding assays; quantitative modelling (led by Dr G Civelekoglu-Scholey); and live cell light microscopy combined with the use of antibody inhibition, mutant analysis and RNA interference to elucidate motor protein/polymer dynamics and function in living C. elegans ciliated neurons, sea urchin blastomeres and Drosophila embryonic cells.


  • MCB 110V iBioseminars in Cell and Molecular Biology (See ASCB iBiology webpage "Flipped Cell Biology" class), (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)


  • Scholey Lab

Honors and Awards

  • 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 (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 Fellowship (Postdoctoral Research on 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 American Society for Cell Biology Fellow.

    Professional Societies

    • American Society for Cell Biology, Emeritus Member and ASCB Fellow.
    • 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 (1993-2017)
    • Editorial Board Member, Molecular Biology of the Cell
    • Biochemical Society (UK) Member
    • NIH Cell Biology Integrated Review Group: Nuclear and Cytoplasmic Structure/Function and Dynamics Study Section (Member 2012 -2015)


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


    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. 

    Khan, S. and Scholey, JM. 2018. Assembly, Functions and Evolution of Archaella, Flagella and Cilia. Current Biology, 28: R278 - R292.