Title(s)Distinguished Professor Emeritus, Molecular and Cellular Biology
SchoolUniversity of California, Davis
Websites
Jonathan M Scholey‬ - ‪Google Scholar
CBS/Molecular & Cell Biology Faculty Profile
Emailjonathanmscholey@gmail.com
jmscholey@ucdavis.edu
 Introduction
Jonathan Scholey is a cell biologist and retired professor with a lifelong interest in the chemistry of cells i.e. the general problem of how atoms and molecules form living, moving, dividing cells. He undertook undergraduate, postgraduate, and postdoctoral studies in cell and molecular biology and 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.

Teaching Interests: Scholey’s teaching reflects his interest in science at the interface of physical and biological sciences and accordingly he has taught various classes in biochemistry, biophysics, cell biology and molecular biology.

Research Focus: Scholey’s research has focused on the cell biology of motor proteins and the mechanisms of mitosis and ciliogenesis.
 Education and Training
Kings College Biophysics Department/MRC Cell Biophysics Unit, London University, London, UKBSc (1st Class Hons)07/1977Cell and Molecular Biology
MRC Laboratory of Molecular Biology and Trinity College, Cambridge University, Cambridge, UKPhD09/1981Molecular Biology
 Awards and Honors
American Society for Cell Biology2017Inducted Fellow
UC Davis2014Distinguished Professor
Fulbright Scholar Program2010  - 2011Fulbright Scholar: Senior Lecturer Award in Molecular Biology, Bosphorus Univ., Istanbul, Turkey
Gordon Research Conference on Motile and Contractile Systems1996Elected Chairperson
March of Dimes Birth Defects Foundation1986  - 1988Basil O'Connor Starter Scholar Research Award
British Medical Research Council and British and American Heart Foundations1982  - 1986Postdoctoral Fellowships to research Mitosis with Prof JR McIntosh in the Dept of MCDB, UC Boulder
Kings College London Biophysics Department1976Sambrooke Exhibition in Natural Sciences
 Brief Research Summary
As a PhD student at the MRC lab of Molecular Biology in Cambridge, Scholey worked with Dr Jake Kendrick-Jones on the biochemistry of myosin motors involved in muscle contraction and cell motility, 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 dividing cells (Scholey et al, 1985). 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. There, 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 the CELL BIOLOGY OF MOTORS INVOLVED IN THE MECHANISMS OF MITOSIS AND CILIOGENESIS (Ou and Scholey, 2022; Khan and Scholey, 2018; Scholey et al, 2016). At UC Davis, the contributions of the Scholey Lab and their collaborators included; (i) the purification and characterization of heterotrimeric kinesin-2 as an anterograde motor required for the assembly of cilia on swimming sea urchin embryos (Cole et al, 1993; Wedaman et al, 1996; Morris and Scholey, 1997); (ii) the finding that kinesin-2 motors exist in both heterotrimeric and homodimeric forms and cooperate with each other and with the retrograde motor, dynein-2, to drive bidirectional intraflagellar transport and ciliogenesis on C. elegans sensory neurons (Signor et al, 1999a,b; Snow, Ou et al, 2004; Pan et al, 2006, Prevo et al, 2015); (iii) the determination of the bipolar homotetrameric organization of mitotic kinesin-5 motors and an analysis of their cooperation with other motors in driving the sliding-filament/force-balance mechanism underlying mitotic spindle assembly and anaphase B spindle elongation in Drosophila embryos (Kashina et al, 1996; Sharp et al, 1999; Brust-Mascher, Civelekoglu-Scholey et al, 2004; Civelekoglu-Scholey et al, 2010; Scholey et al, 2014, 2016); and (iv) the development of a multiple motor-dependent force-balance model for chromosome motility during mitosis in Drosophila embryos (Civelekoglu-Scholey et al, 2006) based on experimental data (Sharp et al, 2000; Rogers et al, 2004).

Scholey also enjoyed teaching various classes in Biochemistry (BIS102, MCB221D); Biophysics (MCB 143); Cell Biology (BIS104, MCB142, MCB110V, MCB212); and Molecular Biology (BIS1A). He retired on his 60th birthday on 03/07/2015 and spends his retirement reading research papers and writing reviews in Cell Biology; reading science books in broad areas of Physics, Chemistry and Biology (his A-level subjects) and in the History of Science (a long-term layperson hobby); following sports, especially the Man United, Fenerbahce and England soccer teams as well as the SF 49ers in the NFL; and learning Turkish and exploring Turkey where he lives in Istanbul with his Turkish Mathematician wife for much of the year.
 Interests
1st Cell Biology & Chemistry; 2nd Molecular Biology; 3rd Physics & Physiology; 4th Zoology & Geology
 Selected Publications
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Altmetrics Details PMC Citations indicate the number of times the publication was cited by articles in PubMed Central (as of September 2022), and the Altmetric score represents citations in news articles and social media. (Note that publications are often cited in additional ways that are not shown here.) Fields are based on how the National Library of Medicine (NLM) classifies the publication's journal and might not represent the specific topic of the publication. Translation tags are based on the publication type and the MeSH terms NLM assigns to the publication. Some publications (especially newer ones and publications not in PubMed) might not yet be assigned Field or Translation tags.) Click a Field or Translation tag to filter the publications.
  1. Motor Cooperation During Mitosis and Ciliogenesis. Annu Rev Cell Dev Biol. 2022 Oct 6;38:49-74. Ou G, Scholey JM. PMID: 35512258.
    View in: PubMed   Fields:    
  2. Assembly, Functions and Evolution of Archaella, Flagella and Cilia. Curr Biol. 2018 03 19; 28(6):R278-R292. Khan S, Scholey JM. PMID: 29558648.
    View in: PubMed   Mentions: 20     Fields:    Translation:AnimalsCells
  3. Intraflagellar transport: mechanisms of motor action, cooperation, and cargo delivery. FEBS J. 2017 09; 284(18):2905-2931. Prevo B, Scholey JM, Peterman EJG. PMID: 28342295; PMCID: PMC5603355.
    View in: PubMed   Mentions: 68     Fields:    Translation:AnimalsCells
  4. Anaphase B. Biology (Basel). 2016 Dec 08; 5(4). Scholey JM, Civelekoglu-Scholey G, Brust-Mascher I. PMID: 27941648; PMCID: PMC5192431.
    View in: PubMed   Mentions: 19  
  5. Functional differentiation of cooperating kinesin-2 motors orchestrates cargo import and transport in C. elegans cilia. Nat Cell Biol. 2015 Dec; 17(12):1536-45. Prevo B, Mangeol P, Oswald F, Scholey JM, Peterman EJ. PMID: 26523365.
    View in: PubMed   Mentions: 67     Fields:    Translation:AnimalsCells
  6. Structural basis for the assembly of the mitotic motor Kinesin-5 into bipolar tetramers. Elife. 2014 Apr 08; 3:e02217. Scholey JE, Nithianantham S, Scholey JM, Al-Bassam J. PMID: 24714498; PMCID: PMC3978770.
    View in: PubMed   Mentions: 35     Fields:    Translation:AnimalsCells
  7. Patronin mediates a switch from kinesin-13-dependent poleward flux to anaphase B spindle elongation. J Cell Biol. 2013 Oct 14; 203(1):35-46. Wang H, Brust-Mascher I, Civelekoglu-Scholey G, Scholey JM. PMID: 24100293; PMCID: PMC3798244.
    View in: PubMed   Mentions: 22     Fields:    Translation:AnimalsCells
  8. Kinesin-2: a family of heterotrimeric and homodimeric motors with diverse intracellular transport functions. Annu Rev Cell Dev Biol. 2013; 29:443-69. Scholey JM. PMID: 23750925.
    View in: PubMed   Mentions: 66     Fields:    Translation:HumansAnimals
  9. The bipolar assembly domain of the mitotic motor kinesin-5. Nat Commun. 2013; 4:1343. 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. PMID: 23299893; PMCID: PMC3562449.
    View in: PubMed   Mentions: 31     Fields:    Translation:AnimalsCells
  10. Intraflagellar transport delivers tubulin isotypes to sensory cilium middle and distal segments. Nat Cell Biol. 2011 Jun 05; 13(7):790-8. Hao L, Thein M, Brust-Mascher I, Civelekoglu-Scholey G, Lu Y, Acar S, Prevo B, Shaham S, Scholey JM. PMID: 21642982; PMCID: PMC3129367.
    View in: PubMed   Mentions: 81     Fields:    Translation:AnimalsCells
  11. Mitotic force generators and chromosome segregation. Cell Mol Life Sci. 2010 Jul; 67(13):2231-50. Civelekoglu-Scholey G, Scholey JM. PMID: 20221784; PMCID: PMC2883081.
    View in: PubMed   Mentions: 37     Fields:    Translation:AnimalsCells
  12. Prometaphase spindle maintenance by an antagonistic motor-dependent force balance made robust by a disassembling lamin-B envelope. J Cell Biol. 2010 Jan 11; 188(1):49-68. Civelekoglu-Scholey G, Tao L, Brust-Mascher I, Wollman R, Scholey JM. PMID: 20065089; PMCID: PMC2812851.
    View in: PubMed   Mentions: 38     Fields:    Translation:AnimalsCells
  13. Control of mitotic spindle length. Annu Rev Cell Dev Biol. 2010; 26:21-57. Goshima G, Scholey JM. PMID: 20604709.
    View in: PubMed   Mentions: 108     Fields:    Translation:AnimalsCells
  14. Kinesin-5-dependent poleward flux and spindle length control in Drosophila embryo mitosis. Mol Biol Cell. 2009 Mar; 20(6):1749-62. Brust-Mascher I, Sommi P, Cheerambathur DK, Scholey JM. PMID: 19158379; PMCID: PMC2655252.
    View in: PubMed   Mentions: 58     Fields:    Translation:AnimalsCells
  15. Quantitative analysis of an anaphase B switch: predicted role for a microtubule catastrophe gradient. J Cell Biol. 2007 Jun 18; 177(6):995-1004. Cheerambathur DK, Civelekoglu-Scholey G, Brust-Mascher I, Sommi P, Mogilner A, Scholey JM. PMID: 17576796; PMCID: PMC2064360.
    View in: PubMed   Mentions: 39     Fields:    Translation:AnimalsCells
  16. The sensory cilia of Caenorhabditis elegans. WormBook. 2007 Mar 08; 1-22. Inglis PN, Ou G, Leroux MR, Scholey JM. PMID: 18050505.
    View in: PubMed   Mentions: 94     Fields:    Translation:AnimalsCells
  17. Mechanism of transport of IFT particles in C. elegans cilia by the concerted action of kinesin-II and OSM-3 motors. J Cell Biol. 2006 Sep 25; 174(7):1035-45. Pan X, Ou G, Civelekoglu-Scholey G, Blacque OE, Endres NF, Tao L, Mogilner A, Leroux MR, Vale RD, Scholey JM. PMID: 17000880; PMCID: PMC2064394.
    View in: PubMed   Mentions: 98     Fields:    Translation:AnimalsCells
  18. Model of chromosome motility in Drosophila embryos: adaptation of a general mechanism for rapid mitosis. Biophys J. 2006 Jun 01; 90(11):3966-82. Civelekoglu-Scholey G, Sharp DJ, Mogilner A, Scholey JM. PMID: 16533843; PMCID: PMC1459506.
    View in: PubMed   Mentions: 64     Fields:    Translation:AnimalsCells
  19. Functional modulation of IFT kinesins extends the sensory repertoire of ciliated neurons in Caenorhabditis elegans. J Cell Biol. 2006 Feb 27; 172(5):663-9. Evans JE, Snow JJ, Gunnarson AL, Ou G, Stahlberg H, McDonald KL, Scholey JM. PMID: 16492809; PMCID: PMC2063699.
    View in: PubMed   Mentions: 64     Fields:    Translation:AnimalsCells
  20. Functional coordination of intraflagellar transport motors. Nature. 2005 Jul 28; 436(7050):583-7. Ou G, Blacque OE, Snow JJ, Leroux MR, Scholey JM. PMID: 16049494.
    View in: PubMed   Mentions: 185     Fields:    Translation:AnimalsCells
  21. 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. 2004 Nov 09; 101(45):15938-43. Brust-Mascher I, Civelekoglu-Scholey G, Kwon M, Mogilner A, Scholey JM. PMID: 15522967; PMCID: PMC524698.
    View in: PubMed   Mentions: 75     Fields:    Translation:AnimalsCells
  22. Two anterograde intraflagellar transport motors cooperate to build sensory cilia on C. elegans neurons. Nat Cell Biol. 2004 Nov; 6(11):1109-13. Snow JJ, Ou G, Gunnarson AL, Walker MR, Zhou HM, Brust-Mascher I, Scholey JM. PMID: 15489852.
    View in: PubMed   Mentions: 204     Fields:    Translation:AnimalsCells
  23. Two mitotic kinesins cooperate to drive sister chromatid separation during anaphase. Nature. 2004 Jan 22; 427(6972):364-70. Rogers GC, Rogers SL, Schwimmer TA, Ems-McClung SC, Walczak CE, Vale RD, Scholey JM, Sharp DJ. PMID: 14681690.
    View in: PubMed   Mentions: 159     Fields:    Translation:AnimalsCells
  24. The chromokinesin, KLP3A, dives mitotic spindle pole separation during prometaphase and anaphase and facilitates chromatid motility. Mol Biol Cell. 2004 Jan; 15(1):219-33. Kwon M, Morales-Mulia S, Brust-Mascher I, Rogers GC, Sharp DJ, Scholey JM. PMID: 14528012; PMCID: PMC307542.
    View in: PubMed   Mentions: 33     Fields:    Translation:AnimalsCells
  25. Cell division. Nature. 2003 Apr 17; 422(6933):746-52. Scholey JM, Brust-Mascher I, Mogilner A. PMID: 12700768.
    View in: PubMed   Mentions: 110     Fields:    Translation:AnimalsCells
  26. A force balance model of early spindle pole separation in Drosophila embryos. Biophys J. 2003 Feb; 84(2 Pt 1):757-69. Cytrynbaum EN, Scholey JM, Mogilner A. PMID: 12547760; PMCID: PMC1302656.
    View in: PubMed   Mentions: 38     Fields:    Translation:AnimalsCells
  27. Intraflagellar transport. Annu Rev Cell Dev Biol. 2003; 19:423-43. Scholey JM. PMID: 14570576.
    View in: PubMed   Mentions: 185     Fields:    Translation:HumansAnimalsCells
  28. Microtubule flux and sliding in mitotic spindles of Drosophila embryos. Mol Biol Cell. 2002 Nov; 13(11):3967-75. Brust-Mascher I, Scholey JM. PMID: 12429839; PMCID: PMC133607.
    View in: PubMed   Mentions: 46     Fields:    Translation:AnimalsCells
  29. Cytoplasmic dynein is required for poleward chromosome movement during mitosis in Drosophila embryos. Nat Cell Biol. 2000 Dec; 2(12):922-30. Sharp DJ, Rogers GC, Scholey JM. PMID: 11146657.
    View in: PubMed   Mentions: 101     Fields:    Translation:HumansAnimalsCells
  30. Microtubule motors in mitosis. Nature. 2000 Sep 07; 407(6800):41-7. Sharp DJ, Rogers GC, Scholey JM. PMID: 10993066.
    View in: PubMed   Mentions: 201     Fields:    Translation:HumansAnimalsCells
  31. Functional coordination of three mitotic motors in Drosophila embryos. Mol Biol Cell. 2000 Jan; 11(1):241-53. Sharp DJ, Brown HM, Kwon M, Rogers GC, Holland G, Scholey JM. PMID: 10637305; PMCID: PMC14771.
    View in: PubMed   Mentions: 107     Fields:    Translation:HumansAnimalsCells
  32. 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. 1999 Nov 01; 147(3):519-30. Signor D, Wedaman KP, Orozco JT, Dwyer ND, Bargmann CI, Rose LS, Scholey JM. PMID: 10545497; PMCID: PMC2151193.
    View in: PubMed   Mentions: 128     Fields:    Translation:AnimalsCells
  33. Antagonistic microtubule-sliding motors position mitotic centrosomes in Drosophila early embryos. Nat Cell Biol. 1999 May; 1(1):51-4. Sharp DJ, Yu KR, Sisson JC, Sullivan W, Scholey JM. PMID: 10559864.
    View in: PubMed   Mentions: 104     Fields:    Translation:AnimalsCells
  34. Two heteromeric kinesin complexes in chemosensory neurons and sensory cilia of Caenorhabditis elegans. Mol Biol Cell. 1999 Feb; 10(2):345-60. Signor D, Wedaman KP, Rose LS, Scholey JM. PMID: 9950681; PMCID: PMC25173.
    View in: PubMed   Mentions: 60     Fields:    Translation:AnimalsCells
  35. The bipolar kinesin, KLP61F, cross-links microtubules within interpolar microtubule bundles of Drosophila embryonic mitotic spindles. J Cell Biol. 1999 Jan 11; 144(1):125-38. Sharp DJ, McDonald KL, Brown HM, Matthies HJ, Walczak C, Vale RD, Mitchison TJ, Scholey JM. PMID: 9885249; PMCID: PMC2148119.
    View in: PubMed   Mentions: 143     Fields:    Translation:AnimalsCells
  36. Heterotrimeric kinesin-II is required for the assembly of motile 9+2 ciliary axonemes on sea urchin embryos. J Cell Biol. 1997 Sep 08; 138(5):1009-22. Morris RL, Scholey JM. PMID: 9281580; PMCID: PMC2136763.
    View in: PubMed   Mentions: 70     Fields:    Translation:AnimalsCells
  37. Kinesin-II, a membrane traffic motor in axons, axonemes, and spindles. J Cell Biol. 1996 Apr; 133(1):1-4. Scholey JM. PMID: 8601599; PMCID: PMC2120781.
    View in: PubMed   Mentions: 37     Fields:    Translation:AnimalsCells
  38. Sequence and submolecular localization of the 115-kD accessory subunit of the heterotrimeric kinesin-II (KRP85/95) complex. J Cell Biol. 1996 Feb; 132(3):371-80. Wedaman KP, Meyer DW, Rashid DJ, Cole DG, Scholey JM. PMID: 8636215; PMCID: PMC2120715.
    View in: PubMed   Mentions: 61     Fields:    Translation:AnimalsCells
  39. A bipolar kinesin. Nature. 1996 Jan 18; 379(6562):270-2. Kashina AS, Baskin RJ, Cole DG, Wedaman KP, Saxton WM, Scholey JM. PMID: 8538794; PMCID: PMC3203953.
    View in: PubMed   Mentions: 165     Fields:    Translation:AnimalsCells
  40. A "slow" homotetrameric kinesin-related motor protein purified from Drosophila embryos. J Biol Chem. 1994 Sep 16; 269(37):22913-6. Cole DG, Saxton WM, Sheehan KB, Scholey JM. PMID: 8083185; PMCID: PMC3201834.
    View in: PubMed   Mentions: 78     Fields:    Translation:Animals
  41. The carboxyl-terminal domain of kinesin heavy chain is important for membrane binding. J Biol Chem. 1994 Jan 14; 269(2):1477-85. Skoufias DA, Cole DG, Wedaman KP, Scholey JM. PMID: 8288613.
    View in: PubMed   Mentions: 40     Fields:    Translation:AnimalsCells
  42. Novel heterotrimeric kinesin-related protein purified from sea urchin eggs. Nature. 1993 Nov 18; 366(6452):268-70. Cole DG, Chinn SW, Wedaman KP, Hall K, Vuong T, Scholey JM. PMID: 8232586.
    View in: PubMed   Mentions: 123     Fields:    Translation:AnimalsCells
  43. Roles of kinesin and kinesin-like proteins in sea urchin embryonic cell division: evaluation using antibody microinjection. J Cell Biol. 1993 Nov; 123(3):681-9. Wright BD, Terasaki M, Scholey JM. PMID: 8227132; PMCID: PMC2200125.
    View in: PubMed   Mentions: 31     Fields:    Translation:AnimalsCells
  44. Isolation of a sea urchin egg kinesin-related protein using peptide antibodies. J Cell Sci. 1992 Feb; 101 ( Pt 2):291-301. Cole DG, Cande WZ, Baskin RJ, Skoufias DA, Hogan CJ, Scholey JM. PMID: 1629246.
    View in: PubMed   Mentions: 52     Fields:    Translation:AnimalsCells
  45. 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. 1991 May; 113(4):817-33. Wright BD, Henson JH, Wedaman KP, Willy PJ, Morand JN, Scholey JM. PMID: 1827446; PMCID: PMC2288992.
    View in: PubMed   Mentions: 38     Fields:    Translation:AnimalsCells
  46. Identification of globular mechanochemical heads of kinesin. Nature. 1989 Mar 23; 338(6213):355-7. Scholey JM, Heuser J, Yang JT, Goldstein LS. PMID: 2493586.
    View in: PubMed   Mentions: 64     Fields:    Translation:AnimalsCells
  47. Inhibition of kinesin-driven microtubule motility by monoclonal antibodies to kinesin heavy chains. J Cell Biol. 1988 Dec; 107(6 Pt 2):2657-67. Ingold AL, Cohn SA, Scholey JM. PMID: 2974459; PMCID: PMC2115674.
    View in: PubMed   Mentions: 66     Fields:    Translation:AnimalsCells
  48. Identification of kinesin in sea urchin eggs, and evidence for its localization in the mitotic spindle. Nature. 1985 Dec 5-11; 318(6045):483-6. Scholey JM, Porter ME, Grissom PM, McIntosh JR. PMID: 2933590.
    View in: PubMed   Mentions: 77     Fields:    Translation:AnimalsCells
  49. Studies on the effect of phosphorylation of the 20,000 Mr light chain of vertebrate smooth muscle myosin. J Mol Biol. 1983 Mar 25; 165(1):139-62. Kendrick-Jones J, Cande WZ, Tooth PJ, Smith RC, Scholey JM. PMID: 6133003.
    View in: PubMed   Mentions: 23     Fields:    Translation:AnimalsCells
  50. Regulation of non-muscle myosin assembly by calmodulin-dependent light chain kinase. Nature. 1980 Sep 18; 287(5779):233-5. Scholey JM, Taylor KA, Kendrick-Jones J. PMID: 6893621.
    View in: PubMed   Mentions: 105     Fields:    Translation:AnimalsCells
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