Jonathan M Scholey – Cell Biologist

TitleDistinguished Professor Emeritus
University/DepartmentMolecular and Cell Biology, University of California, Davis
Websites Jonathan M Scholey‬ – ‪Google Scholar
CBS/Molecular & Cell Biology Faculty Profile


Jonathan Scholey is a cell biologist and retired professor with a lifelong interest in the chemistry of cells i.e. how atoms and molecules form living, moving, dividing cells. After completing A-levels in chemistry, physics (with maths) & biology at grammar school, followed by a gap year laboring on building sites and traveling, Scholey undertook undergraduate, postgraduate, and postdoctoral studies in cell and molecular biology. He was subsequently appointed a faculty member, first in the division of molecular and cell biology at National Jewish Hospital and Research Center in Denver, then at the University of California, Davis, from which he retired on his 60th birthday in 2015. He and his mathematician wife, Gül, currently split time between homes in Davis and Istanbul (where they co-taught a quantitative cell biology class) and enjoy visiting sites of biological, geological and archaeological interest. In 2017 he had successful pituitary surgery at UCD Med Center for the rare neuroendocrine disease, acromegaly, and in 2023 had successful bilateral knee replacement surgery in Istanbul.


Kings College Biophysics Department/MRC Cell Biophysics Unit, London University, London, UKBSc (1st class honours)07/1977Cell and Molecular Biology
MRC Laboratory of Molecular Biology and Trinity College, Cambridge University, Cambridge, UKPhD09/1981Molecular Biology


Scientific Interests: 1. Cell Biology & Chemistry; 2. Molecular Biology; 3. Physics & Physiology; 4. Zoology, Geology & Evolutionary Biology; 5. History of Science.

Teaching: Scholey’s teaching reflects his interest in science at the interface of physical and biological sciences and accordingly he taught various classes at UCD in biochemistry (BIS102, MCB221D), biophysics (MCB143), cell biology (BIS104, MCB142, MCB110V, MCB212) and molecular biology (BIS1A). He taught similar classes during 2 sabbaticals at Bosphorus University in Istanbul. Previously he had tutored undergrad biochemistry at Trinity College, Cambridge and lectured in bioenergetics and the cytoskeleton to grad students at CUHSC, Denver.

Research: Scholey’s research focused on the cell biology of motor proteins and the mechanisms of mitosis and ciliogenesis (see Research Summary below and reviews in Annu. Rev. Cell Dev. Biol. by Scholey 2003; Goshima and Scholey, 2010; Scholey, 2013; Ou and Scholey, 2022).

Quotes he likes: [1] Almost all aspects of life are engineered at the molecular level, and without understanding molecules we can only have a very sketchy understanding of life itself. All approaches at higher levels are suspect until confirmed at the molecular level – Francis Crick. [2] It was obvious – to me at any rate – how an enzyme is able to speed up a chemical reaction by as much as 10 million times. It had to do so by lowering the energy of activation – the energy of forming the activated complex. It could do this by forming strong bonds with the activated complex but only weak bonds with the reactants and products – Linus Pauling. [3] I know I cannot explain the mechanism yet, but the sliding of actin and myosin filaments is a fact Jean Hanson. [4] Man, like other organisms, is so perfectly coordinated that he may easily forget, whether awake or asleep, that he is a colony of cells in action, and that it is the cells which achieve, through him, what he has the illusion of accomplishing himself –Albert Claude. [5] There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved – Charles Darwin. [6] It will be possible, through the detailed determination of amino-acid sequences of hemoglobin molecules and of other molecules too, to obtain much information about the course of the evolutionary process, and to illuminate the question of the origin of species – Linus Pauling. [7] The more clearly we can focus our attention on the wonders and realities of the universe about us, the less taste we shall have for destruction – Rachel Carson.

Current Biology (2006) Q and A interview:

Honorific Awards and Service

American Society for Cell Biology2017Inducted Fellow
UC Davis2014Distinguished Professor
Students of the UCD Biochemistry, Molecular, Cell and Developmental Biology Graduate Group2014Lifetime Achievement Awardee
NIH Cell Biology (NCSD) study section2012-2015Member
Fulbright Scholar Program2010-2011 Fulbright Scholar: Senior Lecturer Award in Molecular Biology, Bosphorus Univ., Istanbul, Turkey
American Society for Cell Biology and Biochemical Society (UK)CurrentEmeritus Member
Sigma Xi; Richard Dawkins Foundation/CFI; AAASCurrentMember
1. Journal of Biological Chemistry; 2. Molecular Biology of the Cell; and 3. CytoskeletonVarious datesEditorial Board Member
Gordon Research Conference on Motile and Contractile Systems1996Elected Chairperson
March of Dimes Birth Defects Foundation1986-1988 Basil O’Connor Starter Scholar Research Awardee
British Medical Research Council and British and American Heart Foundations1982-1986 Postdoctoral Fellowships to research Mitosis with Prof JR McIntosh in the Dept of MCDB, UC Boulder
Kings College London Biophysics Department1976Sambrooke Exhibition in Natural Sciences

Research Summary

As a PhD student at the MRC laboratory of Molecular Biology in Cambridge, Scholey worked with Dr Jake Kendrick-Jones on the biochemistry of myosin-2 motors involved in smooth muscle contraction, non-muscle cell motility, and cytokinesis, focusing on their regulation by Ca++ ions and light chain phosphorylation (Scholey et al, 1980; Kendrick-Jones et al, 1983). Subsequently, between 1982-1986, he undertook postdoctoral studies on mitotic motors (kinesins and dyneins) in the cell biology laboratory of Dr Dick McIntosh at the Dept of MCDB, University of Colorado, Boulder. It was found that kinesin motors, first discovered as axonal transport motors, localize to mitotic spindles in some dividing cells (Scholey et al, 1985; Porter et al, 1987). This work provided the foundation for the future research of the Scholey laboratory, initially during 1986-1989 in the Division of Molecular and Cell Biology at National Jewish Hospital and Research Center in Denver, where monoclonal antibodies that inhibit kinesin-driven motility were developed and were used to help identify the mechanochemical “motor domains” of kinesin motors (Ingold et al, 1988; Scholey et al, 1989).

Scholey moved his lab from NJHRC in Denver to join the faculty of the Depts of Zoology and Molecular and Cellular Biology at UC Davis in 1989. His research at UC Davis was focused on Mitotic motors, IFT motors, and the mechanisms of mitosis and ciliogenesis (Reviews: Sharp et al, 2000a; Inglis et al, 2007; Prevo et al, 2017; Khan and Scholey, 2018; Scholey et al, 2003; 2016). At UC Davis, the contributions of the Scholey Lab and their collaborators included;

[A] Ciliogenesis: (i) the discovery, purification and characterization of heterotrimeric kinesin-2 as an anterograde transport motor required for the assembly of cilia on swimming sea urchin embryos (Cole et al, 1993; Wedaman et al, 1996; Morris and Scholey, 1997) which led the lab into the newly established field of intraflagellar transport (IFT); (ii) the finding that kinesin-2 motors exist in both heterotrimeric and homodimeric forms within sensory cilia on C. elegans neurons (Signor et al, 1999a); (iii) the development and application of IFT assays using fluorescent GFP-reporters in living cells (Orozco et al, 1999) to illuminate how these two forms of kinesin-2 cooperate to drive sequential pathways of anterograde IFT that deliver tubulins to build two distinct ciliary domains (the proximal axoneme core & a distal signaling domain) and, with the retrograde motor, dynein-2, mediate bidirectional IFT and ciliogenesis (Signor et al, 1999b; Snow, Ou et al, 2004; Ou et al, 2005; Evans et al, 2006; Pan et al, 2006, Hao et al, 2011; Prevo et al, 2015).

[B] Mitosis: (i) The 1st purification of the native kinesin-5 mitotic motor (done using Drosophila embryos) as a bipolar homotetramer capable of crosslinking adjacent MTs to drive a “sliding filament mechanism” (Cole et al, 1994; Kashina et al, 1996a,b; Sharp et al, 1999a; Van den Wildenberg et al, 2008; Acar et al, 2013; Scholey et al, 2014); (ii) an analysis of kinesin-5’s cooperation with other motors (dynein; kinesins-4,-8,-13,-14), MT crosslinkers (Ase1/Feo) and MT polymer dynamics to drive the sliding-filament/force-balance mechanism that contributes to mitotic spindle assembly and length control, metaphase poleward flux and anaphase B spindle elongation in Drosophila embryos (Sharp et al, 1999b, 2000c; Brust-Mascher et al, 2004, 2009; Cheerambathur et al, 2007; Wang et al, 2013, 2015; Scholey et al, 2016); (iii) the development of a multiple motor-dependent force-balance model for chromosome motility during the atypically fast mitoses of Drosophila embryos (Civelekoglu-Scholey et al, 2006) based on experimental data (Sharp et al, 2000b; Rogers et al, 2004); (iv) the proposal that kinesin-1-bound mitotic spindle membranes in sea urchin embryos provide exocytotic vesicles for delivery to the cell surface to maintain and repair the plasma membrane, rather than being required for mitosis (Wright et al, 1991, 1993; Bi et al, 1997) whereas in Drosophila embryos, nuclear envelope-derived spindle membranes cooperate with mitotic motors to stabilize the prometaphase spindle as chromosomes are captured (Civelekoglu-Scholey et al, 2010).

2015 Retirement Research Seminar:


Acar S, Carlson DB, Budamagunta MS, Yarov-Yarovoy V, Correia JJ, Niñonuevo MR, Jia W, Tao L, Leary JA, Voss JC, Evans JE, Scholey JM (2013). The bipolar assembly domain of the mitotic motor kinesin-5. Nat Commun.; 4:1343. PMID: 23299893; PMCID: PMC3562449.
Bi GQ, Morris RL, Liao G, Alderton JM, ScholeyJM, Steinhardt RA (1997). Kinesin- and myosin-driven steps of vesicle recruitment for Ca2+-regulated exocytosis. J Cell Biol.;138(5):999-1008. doi: 10.1083/jcb.138.5.999. PMID: 9281579; PMCID: PMC2136755.
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 U S A; 101(45):15938-43. PMID: 15522967; PMCID: PMC524698.
Brust-Mascher I, Sommi P, Cheerambathur DK, Scholey JM (2009). Kinesin-5-dependent poleward flux and spindle length control in Drosophila embryo mitosis. Mol Biol Cell.; 20(6):1749-62. PMID: 19158379; PMCID: PMC2655252.
Cheerambathur DK, Civelekoglu-Scholey G, Brust-Mascher I, Sommi P, Mogilner A, Scholey JM (2007). Quantitative analysis of an anaphase B switch: predicted role for a microtubule catastrophe gradient. J Cell Biol.; 177(6):995-1004. PMID: 17576796; PMCID: PMC2064360.
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. Biophys J.; 90(11):3966-82. PMID: 16533843; PMCID: PMC1459506.
Civelekoglu-Scholey G, Tao L, Brust-Mascher I, Wollman R, Scholey JM 2010). Prometaphase spindle maintenance by an antagonistic motor-dependent force balance made robust by a disassembling lamin-B envelope. J Cell Biol.; 188(1):49-68. PMID: 20065089; PMCID: PMC2812851.
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(6452):268-70. PMID: 8232586.
Cole DG, Saxton WM, Sheehan KB, Scholey JM (1994). A “slow” homotetrameric kinesin-related motor protein purified from Drosophila embryos. J Biol Chem.; 269(37):22913-6. PMID: 8083185; PMCID: PMC3201834.
Evans JE, Snow JJ, Gunnarson AL, Ou G, Stahlberg H, McDonald KL, Scholey JM (2006). Functional modulation of IFT kinesins extends the sensory repertoire of ciliated neurons in Caenorhabditis elegans. J Cell Biol.; 172(5):663-9. PMID: 16492809; PMCID: PMC2063699.
Goshima G, Scholey JM (2010). Control of mitotic spindle length. Annu Rev Cell Dev Biol.; 26:21-57. PMID: 20604709.
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. Nat Cell Biol.;13(7):790-8. PMID: 21642982; PMCID: PMC3129367.
Inglis PN, Ou G, Leroux MR, Scholey JM (2007). The sensory cilia of Caenorhabditis elegans. WormBook; 1-22. doi: 10.1895/wormbook.1.126.2. PMID: 18050505; PMCID: PMC10083726.
Ingold AL, Cohn SA, Scholey JM (1988). Inhibition of kinesin-driven microtubule motility by monoclonal antibodies to kinesin heavy chains. J Cell Biol.; 107(6 Pt 2):2657-67. PMID: 2974459; PMCID: PMC2115674.
Kashina AS, Baskin RJ, Cole DG, Wedaman KP, Saxton WM, Scholey JM (1996a). A bipolar kinesin. Nature; 379(6562):270-2. PMID: 8538794; PMCID: PMC3203953.
Kashina AS, Scholey JM, Leszyk JD, Saxton WM (1996b). An essential bipolar mitotic motor. Nature;384(6606):225. PMID: 8918872; PMCID: PMC3209954.
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. J Mol Biol.; 165(1):139-62. PMID: 6133003.
Khan S, Scholey JM (2018). Assembly, Functions and Evolution of Archaella, Flagella and Cilia. Curr Biol; 28(6):R278-R292. PMID: 29558648.
Morris RL, Scholey JM (1997). Heterotrimeric kinesin-II is required for the assembly of motile 9+2 ciliary axonemes on sea urchin embryos. J Cell Biol.; 138(5):1009-22. PMID: 9281580; PMCID: PMC2136763.
Orozco JT, Wedaman KP, Signor D, Brown H, Rose L, Scholey JM (1999). Movement of motor and cargo along cilia. Nature; 398(6729):674. doi: 10.1038/19448. PMID: 10227290.
Ou G, Blacque OE, Snow JJ, Leroux MR, Scholey JM (2005). Functional coordination of intraflagellar transport motors. Nature; 436(7050):583-7. PMID: 16049494.
Ou G, Scholey JM (2022). Motor Cooperation During Mitosis and Ciliogenesis. Annu Rev Cell Dev Biol.; 38:49-74. PMID: 35512258.
Pan X, Ou G, Civelekoglu-Scholey G, Blacque OE, Endres NF, Tao L, Mogilner A, Leroux MR, Vale RD, Scholey JM (2006). Mechanism of transport of IFT particles in C. elegans cilia by the concerted action of kinesin-II and OSM-3 motors. J Cell Biol.; 174(7):1035-45. PMID: 17000880; PMCID: PMC2064394.
Porter ME, Scholey JM, Stemple DL, Vigers GP, Vale RD, Sheetz MP, McIntosh JR (1987). Characterization of the microtubule movement produced by sea urchin egg kinesin. J Biol Chem.; 262(6):2794-802. PMID: 3102475.
Prevo B, Mangeol P, Oswald F, Scholey JM, Peterman EJ (2015). Functional differentiation of cooperating kinesin-2 motors orchestrates cargo import and transport in C. elegans cilia. Nat Cell Biol. 2015; 17(12):1536-45. PMID: 26523365.
Prevo B, Scholey JM, Peterman EJG (2017). Intraflagellar transport: mechanisms of motor action, cooperation, and cargo delivery. FEBS J.; 284(18):2905-2931. PMID: 28342295; PMCID: PMC5603355.
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(6972):364-70. PMID: 14681690.
Scholey JM (2003). Intraflagellar transport. Annu Rev Cell Dev Biol.; 19:423-43. PMID: 14570576.
Scholey JM (2013). Kinesin-2: a family of heterotrimeric and homodimeric motors with diverse intracellular transport functions. Annu Rev Cell Dev Biol.; 29:443-69. PMID: 23750925.
Scholey JM, Brust-Mascher I, Mogilner A (2003). Cell division. Nature; 422(6933):746-52. PMID: 12700768.
Scholey JM, Civelekoglu-Scholey G, Brust-Mascher I (2016). Anaphase B. Biology (Basel). 5(4). PMID: 27941648; PMCID: PMC5192431.
Scholey JM, Heuser J, Yang JT, Goldstein LS (1989). Identification of globular mechanochemical heads of kinesin. Nature; 338(6213):355-7. PMID: 2493586.
Scholey JE, Nithianantham S, Scholey JM, Al-Bassam J (2014). Structural basis for the assembly of the mitotic motor Kinesin-5 into bipolar tetramers. Elife.; 3:e02217. PMID: 24714498; PMCID: PMC3978770.
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(6045):483-6. PMID: 2933590.
Scholey JM, Taylor KA, Kendrick-Jones (1980). Regulation of non-muscle myosin assembly by calmodulin-dependent light chain kinase. Nature; 287(5779):233-5. J. PMID: 6893621.
Sharp DJ, McDonald KL, Brown HM, Matthies HJ, Walczak C, Vale RD, Mitchison TJ, Scholey JM (1999a). The bipolar kinesin, KLP61F, cross-links microtubules within interpolar microtubule bundles of Drosophila embryonic mitotic spindles. J Cell Biol. 1999a; 144(1):125-38. PMID: 9885249; PMCID: PMC2148119.
Sharp DJ, Yu KR, Sisson JC, Sullivan W, Scholey JM (1999b). Antagonistic microtubule-sliding motors position mitotic centrosomes in Drosophila early embryos. Nat Cell Biol.; 1(1):51-4. PMID: 10559864.
Sharp DJ, Rogers GC, Scholey JM (2000a). Microtubule motors in mitosis. Nature; 407(6800):41-7. PMID: 10993066.
Sharp DJ, Rogers GC, Scholey JM (2000b). Cytoplasmic dynein is required for poleward chromosome movement during mitosis in Drosophila embryos. Nat Cell Biol.; 2(12):922-30. PMID: 11146657.
Sharp DJ, Brown HM, Kwon M, Rogers GC, Holland G, Scholey JM (2000c). Functional coordination of three mitotic motors in Drosophila embryos. Mol Biol Cell. 11(1):241-53. PMID: 10637305; PMCID: PMC14771.
Signor D, Wedaman KP, Rose LS, Scholey JM (1999a). Two heteromeric kinesin complexes in chemosensory neurons and sensory cilia of Caenorhabditis elegans. Mol Biol Cell.; 10(2):345-60. PMID: 9950681; PMCID: PMC25173.
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. J Cell Biol.; 147(3):519-30. PMID: 10545497; PMCID: PMC2151193.
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. Nat Cell Biol.; 6(11):1109-13. PMID: 15489852.
van den Wildenberg SM, Tao L, Kapitein LC, Schmidt CF, Scholey JM, Peterman EJ (2008). The homotetrameric kinesin-5 KLP61F preferentially crosslinks microtubules into antiparallel orientations. Curr Biol.; 18(23):1860-4. PMID: 19062285; PMCID: PMC2657206.
Wang H, Brust-Mascher I, Scholey JM (2015). The microtubule cross-linker Feo controls the midzone stability, motor composition, and elongation of the anaphase B spindle in Drosophila embryos. Mol Biol Cell.;26(8):1452-62. PMID: 25694445; PMCID: PMC4395126.
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. J Cell Biol.; 203(1):35-46. PMID: 24100293; PMCID: PMC3798244.
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. J Cell Biol.; 132(3):371-80. PMID: 8636215; PMCID: PMC2120715.
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. J Cell Biol.; 113(4):817-33. PMID: 1827446; PMCID: PMC2288992.
Wright BD, Terasaki M, Scholey JM (1993). Roles of kinesin and kinesin-like proteins in sea urchin embryonic cell division: evaluation using antibody microinjection. J Cell Biol.; 123(3):681-9. PMID: 8227132; PMCID: PMC2200125.