Jonathan M. Scholey

Jonathan Scholey

Position Title
Distinguished Professor Emeritus

  • Department of Molecular and Cellular Biology, College of Biological Sciences
3253 Green Hall

Profile Introduction

My research has focused on the Cell Biology of Motor Proteins and the Mechanisms of Mitosis and Ciliogenesis. I studied myosin motors as a PhD student with Dr Jake Kendrick-Jones at the MRC Laboratory of Molecular Biology in Cambridge and mitotic motors (kinesins and dyneins) as a post-doc with Dr Dick McIntosh in MCDB at CU Boulder. Being a member of those two labs provided an excellent foundation for my own 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. This began at grammar school in York (UK) and, after I moved to Northampton grammar school, a very formal institution to which I was not well-suited, I maintained enough interest in science to complete A-levels in (i) biology (ii) chemistry and (iii) physics with supplemental maths and statistics (also earning an S (scholarship)-level in chemistry). Thankfully, following a year outside academia sampling life as a laborer on building sites, this got me a (pre-Thatcher era) government grant to study science at university. Consequently, after obtaining a BSc (1st class) in cell and molecular biology from the excellent Biophysics Dept at KCL; a PhD in molecular biology from the MRC Laboratory of Molecular Biology and Trinity College, Cambridge (where I periodically tutored undergraduate biochemistry); and then doing post-doctoral and junior faculty research in Colorado (USA), I was privileged and happy to teach various classes in biochemistry, biophysics, cell biology and molecular biology as a professor at UC Davis (and, during sabbaticals, at Bosphorus University in Istanbul). I retired on my 60th birthday and now enjoy following science as a hobby.

Research Interests

Cell Biology: Motor Proteins in Mitosis and Ciliogenesis.

Specific Projects: 1. Mechanisms of mitotic spindle assembly, anaphase B spindle elongation and chromosome segregation in Drosophila and sea urchin embryos (funded by NIH grant GM55507): 2. Intraflagellar transport (IFT) motors 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, molecular biology and microscopy; motor protein purification and motility/gliding assays; motor gene cloning, sequencing, expression; antibody production/microinjection; 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 dynamics and function in living C. elegans ciliated neurons, sea urchin blastomeres and Drosophila embryos.

The research contributions of the Scholey lab and our collaborators (see selected publications) include;

(i) use of motility-inhibiting monoclonal antibodies to identify globular "motor domains" on kinesin motors (Ingold et al, 1988; Scholey et al, 1989);

(ii) hypothesis that sea urchin embryo mitotic spindles contain kinesin-1-bound vesicles (Wright et al, 1991) that are not required for mitosis/cell division (Wright et al, 1993) but instead move to the cell surface for membrane insertion via exocytosis (Bi et al, 1997);

(iii) discovery of kinesin-2 as a novel, heterotrimeric, anterograde MT motor (Cole et al, 1993; Wedaman et al, 1996) required for ciliogenesis on swimming sea urchin embryos (Morris and Scholey, 1997);

(iv) identification of heterotrimeric and homodimeric forms of kinesin-2 (Signor et al, 1999a) and demonstration of their cooperation, together with retrograde IFT-dynein (Signor et al, 1999b), in IFT and sensory ciliogenesis on C. elegans neurons (Snow, Ou et al, 2004; Ou et al, 2005; Prevo et al, 2015);

(v) determination of bipolar, homotetrameric organization of purified kinesin-5 motors as likely "sliding filament" motors in mitosis  (Cole et al, 1994; Kashina et al, 1996; Acar et al, 2013);

(vi) proposal of a multiple motor-dependent sliding filament/force-balance mechanism for mitosis in Drosophila embryos (Sharp et al, 1999, 2000a, c);

(vii) development of a quantitative sliding filament model for anaphase B in Drosophila embryos based on a switch from poleward flux to spindle elongation (Brust-Mascher and Scholey, 2002; Brust-Mascher, Civelekoglu-Scholey et al, 2004; Cheerambathur et al, 2007; Wang et al, 2013);

(viii) development of quantitative models for motor-dependent chromosome motility (Civelekoglu-Scholey et al, 2006) based on studies of live mitosis (Sharp et al, 2000b; Rogers et al, 2004). 


  • MCB 110V iBioseminars in Cell and Molecular Biology (2012-2015: See ASCB iBiology webpage "Flipped Cell Biology" class)
  • BIO 494 Quantitative Cell and Molecular Biology, (Fall 2013: Bogazici University Molecular Biology and Genetics Major - mathematical modeling of fundamental cell biological topics, co-taught with Dr Gul Civelekoglu-Scholey)
  • MCB 143 Cell and Molecular Biophysics, (2001-2012) - an elective for advanced undergrad students in the cell biology, BMB and genetics majors whose content is outlined in Scholey, 2013, Mol. Biol. Cell., 24; 433-439 
  • MCB 212 Cell Biology, (Winter 2012) - a BMCDB graduate group core course
  • MCB 221D Cellular Biochemistry, (1991-2012) - a BMB graduate group core course
  • BIO 580 Molecular Biology of Mitosis and Cell Division, (Fall semester 2010: Bogazici University Molecular Biology and Genetics Graduate Group - in depth discussion of emerging research on mechanisms of mitosis/cell division)
  • BIO 495 Cell and Molecular Biophysics, (Fall semester 2010: Bogazici University Molecular Biology and Genetics Major, adapted from MCB 143)
  • BIS 102 Biochemistry: Structure and Function of Biomolecules, (1993-2000) - an upper division biological sciences core course
  • BIS 104 Cell Biology, (1998-2000) - an upper division biological sciences core course
  • MCB 142 Advanced Cell Biology, (1990-1993) - an elective class for seniors
  • BIS 1A Introductory Molecular Biology, (1989-1992) - a lower division core course for biological science majors


  • Scholey Lab

Honors and Awards

  • 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 Traveling 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 (sponsored by the Undergrad Students of the Bis102 Biochemistry Class).
  • 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 Students of 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 Inducted as American Society for Cell Biology Fellow.

Professional Societies

  • American Society for Cell Biology, Emeritus Member and ASCB Fellow.
  • Biochemical Society (UK), 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 (1993-2017)
  • Editorial Board Member, Molecular Biology of the Cell (2005-2021)
  • 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.

Selected Publications

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.

Ingold AL, Cohn SA, Scholey JM. 1988. Inhibition of kinesin-driven microtubule motility by monoclonal antibodies to kinesin heavy chain. Journal of Cell Biology, 107: 2657-2667.

Scholey JM, Heuser J, Yang JT, Goldstein LS. 1989. Identification of globular mechanochemical heads of kinesin. Nature, 338: 355-357.

Wright BD, Henson JH, Wedaman KP, Willy PJ, Morand JN, Scholey JM. 1991. Subcellular localization and sequence of sea urchin kinesin heavy chain: Evidence for its association with membranes in the mitotic apparatus and interphase cytoplasm. Journal of Cell Biology, 113: 817-833.

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.

Bi GQ, Morris RL, Liao G, Alderton JM, Scholey JM, Steinhardt RA. 1997. Kinesin- and myosin-driven steps for Ca++-regulated exocytosis. Journal of Cell Biology, 138, 999-1008. 

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.

Signor D, Wedaman KP, Rose LS and Scholey JM. 1999a. Two heteromeric kinesin complexes in chemosensory neurons and sensory cilia of Caenorhabditis elegans. Molecular Biology of the Cell, 10: 345-360.

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. 1999b. 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, Brown HM, Kwon K, Rogers GC, Holland G, Scholey JM. 2000a. Functional coordination of three mitotic motors in Drosophila embryos. Molecular Biology of the Cell. 11: 241-253.

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

Sharp DJ, Rogers GC, Scholey JM. 2000c. Microtubule motors in mitosis. Nature. 407: 41-47.

Brust-Mascher I and Scholey JM. 2002. Microtubule flux and sliding in mitotic spindles of Drosophila embryos. Molecular Biology of the Cell. 13: 3967-3975.

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.

Cheerambathur DK, Civelekoglu-Scholey G, Brust-Mascher I, Sommi P, Mogilner A and Scholey JM. 2007. Quantitative analysis of an anaphase B switch: predicted role for a microtubule catastrophe gradient. Journal of Cell Biology, 177: 995-1004.

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.

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.

Ou, G. and Scholey, JM. 2022. Motor Cooperation during Mitosis and Ciliogenesis. Annual Review of Cell and Developmental Biology. Online ahead of print.