Genome evolution in the allotetraploid frog Xenopus laevis. Session AM, Uno Y, Kwon T, Chapman JA, Toyoda A, Takahashi S, Fukui A, Hikosaka A, Suzuki A, Kondo M, van Heeringen SJ, Quigley I, Heinz S, Ogino H, Ochi H, Hellsten U, Lyons JB, Simakov O, Putnam N, Stites J, Kuroki Y, Tanaka T, Michiue T, Watanabe M, Bogdanovic O, Lister R, Georgiou G, Paranjpe SS, van Kruijsbergen I, Shu S, Carlson J, Kinoshita T, Ohta Y, Mawaribuchi S, Jenkins J, Grimwood J, Schmutz J, Mitros T, Mozaffari SV, Suzuki Y, Haramoto Y, Yamamoto TS, Takagi C, Heald R, Miller K, Haudenschild C, Kitzman J, Nakayama T, Izutsu Y, Robert J, Fortriede J, Burns K, Lotay V, Karimi K, Yasuoka Y, Dichmann DS, Flajnik MF, Houston DW, Shendure J, DuPasquier L, Vize PD, Zorn AM, Ito M, Marcotte EM, Wallingford JB, Ito Y, Asashima M, Ueno N, Matsuda Y, Veenstra GJ, Fujiyama A, Harland RM, Taira M, Rokhsar DS. Nature. 2016 Oct 19;538(7625):336-343. doi: 10.1038/nature19840. PDF

Transcription brings the complex(ity) to the centromere. Grenfell AW, Strzelecka M, Heald R. Cell Cycle. 2016 Oct 13:1-2. [Epub ahead of print] doi: 10.1080/15384101.2016.1242962

Mitotic noncoding RNA processing promotes kinetochore and spindle assembly in Xenopus. Grenfell AW, Heald R, Strzelecka M. J Cell Biol. 2016 Jul 18;214(2):133-41. doi: 10.1083/jcb.201604029. Erratum in: J Cell Biol. 2016 Sep 12;214(6):783. PDF

Microtubules: 50 years on from the discovery of tubulin. Borisy G, Heald R, Howard J, Janke C, Musacchio A, Nogales E. Nat Rev Mol Cell Biol. 2016 Apr 22;17(5):322-8. doi: 10.1038/nrm.2016.45. PDF

A versatile multivariate image analysis pipeline reveals features of Xenopus extract spindles. Grenfell AW, Strzelecka M, Crowder ME, Helmke KJ, Schlaitz AL, Heald R. J Cell Biol. 2016 Apr 11;213(1):127-36. doi: 10.1083/jcb.201509079. PDF


Heald R, Khodjakov A. Thirty years of search and capture: The complex simplicity of mitotic spindle assembly. J Cell Biol. 2015 Dec 21;211(6):1103-11. doi: 10.1083/jcb.201510015. PDF

Levy DL and Heald R. Biological Scaling Problems and Solutions in Amphibians. Cold Spring Harb Perspect Biol. doi: 10.1101/cshperspect.a019166 PDF

Lane AB, Strzelecka M, Ettinger A, Grenfell AW, Wittman T and Heald R. Enzymatically Generated CRISPR Libraries for Genome Labeling and Screening. Dev. Cell 2015 August 10: 34(3):373-378. PubMed PMID: 26212133. PDF

Crowder ME, Strzelecka M, Wilbur JD, Good MC, von Dassow G, and Heald R. A comparative analysis of spindle morphometrics across metazoans. Curr Biol 2015 Jun 1: 25:1-9. PubMed PMID: 26004761. PDF

Miller KE, and Heald R. Glutamylation of Nap1 modulates histone H1 dynamics and chromosome condensation in Xenopus. J Cell Biol 2015 Apr 27: 209(2):211-20. PubMed PMID: 25897082. PDF


Strzelecka M, and Heald R. RUVs drive chromosome decondensation after mitosis. Dev Cell 2014 Nov 10: 31(3):259-60. PubMed PMID: 25453825. PDF

Helmke KJ, and Heald R. TPX2 levels modulate meiotic spindle size and architecture in Xenopus egg extracts. J Cell Biol 2014 Aug 4: 206(3):385-93. PubMed PMID: 25070954. PDF

Heald R, and Cohen-Fix, O. Morphology and function of membrane-bound organelles. Curr Opin Cell Biol 2014 Feb: 26:79-86. PubMed PMID: 24529249. PDF


Good MC, Vahey MD, Skandarajah A, Fletcher DA, Heald R. Cytoplasmic volume modulates spindle size during embryogenesis. Science 2013 Nov 15;342(6160):856-60. PubMed PMID:24233724. PDF

Schlaitz AL, Thompson J, Wong CC, Yates JR 3rd, Heald R. REEP3/4 ensure endoplasmic reticulum clearance from metaphase chromatin and proper nuclear envelope architecture. Dev. Cell 2013 Aug 12;26(3):315-23. PubMed PMID: 23911198l; PubMed Central PMCID: PMC3745822.PDF

Wilbur JD and Heald R. Cryptic No longer: Arrays of CLASP1 TOG Domains. Structure 2013 Jun 4;21(6):869-70. PubMed PMID: 23747108. PDF

Bird SL, Heald R, Weis K. RanGTP and CLASP1 cooperate to position the mitotic spindle. Mol. Biol. Cell Aug;24(16):2506-14. PubMed PMID: 23783028; PubMed Central PMCID:PMC3744954. PDF

Helmke KJ, Heald R, Wilbur JD. Interplay between spindle architecture and function. Int'l Rev.of Cell and Mol. Biol. 2013. 306:83-125. PubMed PMID: 24016524.PDF

Wilbur JD, Heald R. Mitotic spindle scaling during Xenopus development by kif2a and importin α. Elife. 2013; 2:e00290. doi: 10.7554/eLife.00290. Epub 2013 Feb 19. PubMed PMID: 23425906; PubMed Central PMCID: PMC3576809.PDF

Whitehead E, Heald R, Wilbur JD. N-terminal phosphorylation of p60 katanin directly regulates microtubule severing. J Mol. Biol. 2013 Jan 23; 425(2):214-21. PubMed PMID:23178168; PubMed Central PMCID: PMC3540178. PDF


Xiao B, Freedman BS, Miller KE, Jeald R, Marko JF. Histone H1 compact DNA under force and during chromatin assembly. Mol. Biol. Cell. 2012 Dec;23(24):4864-71. PubMed PMID: 23097493; PubMed Central PMID: 23097493. PDF

Levy DL, and Heald R. Mechanisms of intracellular scaling. Annu Rev Cell Dev Biol 2012; 28:113-35. PubMed PMID: 22804576. PDF

Riggs B, Berhman ZJ, Heald R. Altering membrane topology with Sar1 does not impair spindle assembly in Xenopus egg extracts. Cytoskeleton (Hoboken). 2012 Aug;69(8):591-9. PubMed PMID: 22605651. PDF

Patel K, Nogales E, Heald R. Multiple domains of human CLASP contribute to microtubule dynamics and organization in vitro and in Xenopus egg extracts. Cytoskeleton (Hoboken). 2012 Mar; 69(3):155-65. PubMed PMID: 22278908; PubMed Central PMCID: PMC3315288. PDF


Loughlin R, Wilbur JD, McNally FJ, Nédélec FJ, Heald R. Katanin contributes to interspecies spindle length scaling in Xenopus. Cell. 2011 Dec 9;147(6):1397-407. PubMed PMID: 22153081; PubMed Central PMCID: PMC3240848. PDF

Halpin D, Kalab P, Wang J, Weis K, Heald R. Mitotic spindle assembly around RCC1-coated beads in Xenopus egg extracts. PLoS Biol. 2011 Dec;9(12):e1001225. Epub 2011 Dec 27. PubMed PMID: 22215983; PubMed Central PMCID: PMC3246454. PDF

Kieserman EK, Heald R. Mitotic chromosome size scaling in Xenopus. Cell Cycle. 2011 Nov 15;10(22):3863-70. Epub 2011 Nov 15. PubMed PMID: 22071695; PubMed Central PMCID: PMC3266116. PDF

Soderholm JF, Bird SL, Kalab P, Sampathkumar Y, Hasegawa K, Uehara-Bingen M, Weis K, Heald R. Importazole, a small molecule inhibitor of the transport receptor importin-β. ACS Chem Biol. 2011 Jul 15;6(7):700-8. Epub 2011 Apr 21. PubMed PMID: 21469738; PubMed Central PMCID: PMC3137676. PDF

Fu H, Freedman BS, Lim CT, Heald R, Yan J. Atomic force microscope imaging of  chromatin assembled in Xenopus laevis egg extract. Chromosoma. 2011 Jun;120(3):245-54. doi: 10.1007/s00412-010-0307-4. Epub 2011 Jan 11. PubMed PMID: 21369955. PDF

Ridley A, Heald R. Cell structure and dynamics. Curr Opin Cell Biol. 2011 Feb;23(1):1-3. Epub 2010 Dec 27. PubMed PMID: 21190823.


Loughlin R, Heald R, Nédélec F. A computational model predicts Xenopus meiotic spindle organization. J Cell Biol. 2010 Dec 27;191(7):1239-49. Epub 2010 Dec 20.  PubMed PMID: 21173114; PubMed Central PMCID: PMC3010074. PDF

Levy DL, Heald R. Nuclear size is regulated by importin α and Ntf2 in Xenopus. Cell. 2010 Oct 15;143(2):288-98. PubMed PMID: 20946986; PubMed Central PMCID: PMC2966892. PDF

Freedman BS, Miller KE, Heald R. Xenopus egg extracts increase dynamics of histone H1 on sperm chromatin. PLoS One. 2010 Sep 29;5(9). pii: e13111. PubMed PMID: 20927327; PubMed Central PMCID: PMC2947519. PDF

Heald R. Rebecca Heald. Curr Biol. 2010 Jun 22;20(12):R503-4. PubMed PMID: 20627848.

Freedman BS, Heald R. Functional comparison of H1 histones in Xenopus reveals isoform-specific regulation by Cdk1 and RanGTP. Curr Biol. 2010 Jun 8;20(11):1048-52. Epub 2010 May 13. PubMed PMID: 20471264. PDF


Srsen V, Fant X, Heald R, Rabouille C, Merdes A. Centrosome proteins form an  insoluble perinuclear matrix during muscle cell differentiation. BMC Cell Biol. 2009 Apr 21;10:28. PubMed PMID: 19383121.


Heald R. Rebecca Heald: passionate about cycles. Interview by Caitlin Sedwick. J Cell Biol. 2008 Nov 17;183(4):570-1. PubMed PMID: 19015313; PubMed Central PMCID: PMC2582880.

Loughlin R, Riggs B, Heald R. SnapShot: motor proteins in spindle assembly. Cell. 2008 Aug 8;134(3):548-548.e1. PubMed PMID: 18692476.

Kalab P, Heald R. The RanGTP gradient - a GPS for the mitotic spindle. J Cell Sci. 2008 May 15;121(Pt 10):1577-86. PubMed PMID: 18469014.

Andersen CB, Wan Y, Chang JW, Riggs B, Lee C, Liu Y, Sessa F, Villa F, Kwiatkowski N, Suzuki M, Nallan L, Heald R, Musacchio A, Gray NS. Discovery of selective aminothiazole aurora kinase inhibitors. ACS Chem Biol. 2008 Mar 20;3(3):180-92. Epub 2008 Feb 29. PubMed PMID: 18307303.

Walczak CE, Heald R. Mechanisms of mitotic spindle assembly and function. Int Rev Cytol. 2008;265:111-58. Review. PubMed PMID: 18275887.


Blower MD, Feric E, Weis K, Heald R. Genome-wide analysis demonstrates conserved localization of messenger RNAs to mitotic microtubules. J Cell Biol. 2007 Dec 31;179(7):1365-73. PubMed PMID: 18166649; PubMed Central PMCID: PMC2373496.

Heald R. Brinkley-fest of mitosis. Dev Cell. 2007 Aug;13(2):168-76. PubMed PMID: 17681129.

Brown KS, Blower MD, Maresca TJ, Grammer TC, Harland RM, Heald R. Xenopus tropicalis egg extracts provide insight into scaling of the mitotic spindle. J Cell Biol. 2007 Mar 12;176(6):765-70. Epub 2007 Mar 5. PubMed PMID: 17339377; PubMed Central PMCID: PMC2064050.

Yan J, Maresca TJ, Skoko D, Adams CD, Xiao B, Christensen MO, Heald R, Marko  JF. Micromanipulation studies of chromatin fibers in Xenopus egg extracts reveal  ATP-dependent chromatin assembly dynamics. Mol Biol Cell. 2007 Feb;18(2):464-74.  Epub 2006 Nov 15. PubMed PMID: 17108322; PubMed Central PMCID: PMC1783770.


Kenney RD, Heald R. Essential roles for cohesin in kinetochore and spindle function in Xenopus egg extracts. J Cell Sci. 2006 Dec 15;119(Pt 24):5057-66. PubMed PMID: 17158911.

Kaláb P, Pralle A, Isacoff EY, Heald R, Weis K. Analysis of a RanGTP-regulated gradient in mitotic somatic cells. Nature. 2006 Mar 30;440(7084):697-701. PubMed PMID: 16572176.

Maresca TJ, Heald R. The long and the short of it: linker histone H1 is required for metaphase chromosome compaction. Cell Cycle. 2006 Mar;5(6):589-91. Epub 2006 Mar 15. Review. PubMed PMID: 16582611.

Soderholm J, Uehara-Bingen M, Weis K, Heald R. Challenges facing the biologist doing chemical genetics. Nat Chem Biol. 2006 Feb;2(2):55-8. PubMed PMID: 16421579.

Heald R. Cell biology. Serving up a plate of chromosomes. Science. 2006 Jan 20;311(5759):343-4. PubMed PMID: 16424328.

Hannak E, Heald R. Xorbit/CLASP links dynamic microtubules to chromosomes in  the Xenopus meiotic spindle. J Cell Biol. 2006 Jan 2;172(1):19-25. PubMed PMID: 16390996; PubMed Central PMCID: PMC2063525.

Budde PP, Desai A, Heald R. Analysis of microtubule polymerization in vitro and during the cell cycle in Xenopus egg extracts. Methods. 2006 Jan;38(1):29-34. PubMed PMID: 16343934.

Maresca TJ, Heald R. Methods for studying spindle assembly and chromosome condensation in Xenopus egg extracts. Methods Mol Biol. 2006;322:459-74. Review.  PubMed PMID: 16739744.

Hannak E, Heald R. Investigating mitotic spindle assembly and function in vitro using Xenopus laevis egg extracts. Nat Protoc. 2006;1(5):2305-14. PubMed PMID: 17406472.


Maresca TJ, Niederstrasser H, Weis K, Heald R. Xnf7 contributes to spindle integrity through its microtubule-bundling activity. Curr Biol. 2005 Oct 11;15(19):1755-61. PubMed PMID: 16213823.

Maresca TJ, Heald R. Chromosome congression: another fine mesh we've gotten into. Dev Cell. 2005 Sep;9(3):314-5. Review. PubMed PMID: 16139222.

Maresca TJ, Freedman BS, Heald R. Histone H1 is essential for mitotic chromosome architecture and segregation in Xenopus laevis egg extracts. J Cell Biol. 2005 Jun 20;169(6):859-69. PubMed PMID: 15967810; PubMed Central PMCID: PMC2171634.

Blower MD, Nachury M, Heald R, Weis K. A Rae1-containing ribonucleoprotein complex is required for mitotic spindle assembly. Cell. 2005 Apr 22;121(2):223-34. PubMed PMID: 15851029.

Soderholm J, Heald R. Scratch n' screen for inhibitors of cell migration. Chem Biol. 2005 Mar;12(3):263-5. Review. PubMed PMID: 15797209.


Banks JD, Heald R. Adenomatous polyposis coli associates with the microtubule-destabilizing protein XMCAK. Curr Biol. 2004 Nov 23;14(22):2033-8. PubMed PMID: 15556866.

Gadde S, Heald R. Mechanisms and molecules of the mitotic spindle. Curr Biol. 2004 Sep 21;14(18):R797-805. Review. PubMed PMID: 15380094.

Deehan R, Heald R. Centromere glue provides spindle cue. Cell. 2004 Sep 3;118(5):529-30. Review. PubMed PMID: 15339655.

Skop AR, Liu H, Yates J 3rd, Meyer BJ, Heald R. Dissection of the mammalian midbody proteome reveals conserved cytokinesis mechanisms. Science. 2004 Jul 2;305(5680):61-6. Epub 2004 May 27. PubMed PMID: 15166316.

Krauss SW, Lee G, Chasis JA, Mohandas N, Heald R. Two protein 4.1 domains essential for mitotic spindle and aster microtubule dynamics and organization in  vitro. J Biol Chem. 2004 Jun 25;279(26):27591-8. Epub 2004 Apr 21. PubMed PMID: 15102852.

Heald RW. Cell division: burning the spindle at both ends. Nature. 2004 Jan 22;427(6972):300-1. PubMed PMID: 14737150.

Wignall SM, Gray NS, Chang YT, Juarez L, Jacob R, Burlingame A, Schultz PG, Heald R. Identification of a novel protein regulating microtubule stability through a chemical approach. Chem Biol. 2004 Jan;11(1):135-46. PubMed PMID: 15113003.


Wignall SM, Heald R. The role of chromosome architecture in spindle assembly  and anaphase: the condensed version. Cell Cycle. 2003 Nov-Dec;2(6):590-1. PubMed  PMID: 14512775.

Krauss SW, Chen C, Penman S, Heald R. Nuclear actin and protein 4.1: essential interactions during nuclear assembly in vitro. Proc Natl Acad Sci U S A. 2003 Sep 16;100(19):10752-7. Epub 2003 Sep 5. PubMed PMID: 12960380; PubMed Central PMCID: PMC196875.

Wignall SM, Deehan R, Maresca TJ, Heald R. The condensin complex is required  for proper spindle assembly and chromosome segregation in Xenopus egg extracts. J Cell Biol. 2003 Jun 23;161(6):1041-51. PubMed PMID: 12821643; PubMed Central PMCID: PMC2172990.

Budde PP, Heald R. Centrosomes and kinetochores, who needs 'Em? The role of noncentromeric chromatin in spindle assembly. Curr Top Dev Biol. 2003;56:85-113.  Review. PubMed PMID: 14584727.


Krauss SW, Heald R, Lee G, Nunomura W, Gimm JA, Mohandas N, Chasis JA. Two distinct domains of protein 4.1 critical for assembly of functional nuclei in vitro. J Biol Chem. 2002 Nov 15;277(46):44339-46. Epub 2002 Aug 8. PubMed PMID: 12171917.

Kalab P, Weis K, Heald R. Visualization of a Ran-GTP gradient in interphase and mitotic Xenopus egg extracts. Science. 2002 Mar 29;295(5564):2452-6. PubMed PMID: 11923538.

Heald R, Nogales E. Microtubule dynamics. J Cell Sci. 2002 Jan 1;115(Pt 1):3-4. PubMed PMID: 11801717.


Chang YT, Wignall SM, Rosania GR, Gray NS, Hanson SR, Su AI, Merlie J Jr, Moon HS, Sangankar SB, Perez O, Heald R, Schultz PG. Synthesis and biological evaluation of myoseverin derivatives: microtubule assembly inhibitors. J Med Chem. 2001 Dec 20;44(26):4497-500. PubMed PMID: 11741468.

Van Hooser AA, Heald R. Kinetochore function: the complications of becoming attached. Curr Biol. 2001 Oct 30;11(21):R855-7. Review. PubMed PMID: 11696340.

Merlie J Jr, Heald R. Mitotic spindle assembly in vitro. Curr Protoc Cell Biol. 2001 May;Chapter 11:Unit 11.13. PubMed PMID: 18228304.

Budde PP, Kumagai A, Dunphy WG, Heald R. Regulation of Op18 during spindle assembly in Xenopus egg extracts. J Cell Biol. 2001 Apr 2;153(1):149-58. PubMed PMID: 11285281; PubMed Central PMCID: PMC2185528.

Banks JD, Heald R. Chromosome movement: dynein-out at the kinetochore. Curr Biol. 2001 Feb 20;11(4):R128-31. Review. PubMed PMID: 11250166.

Nachury MV, Maresca TJ, Salmon WC, Waterman-Storer CM, Heald R, Weis K. Importin beta is a mitotic target of the small GTPase Ran in spindle assembly. Cell. 2001 Jan 12;104(1):95-106. PubMed PMID: 11163243.

Wignall SM, Heald R. Methods for the study of centrosome-independent spindle  assembly in Xenopus extracts. Methods Cell Biol. 2001;67:241-56. PubMed PMID: 11550472.


Heald R. Motor function in the mitotic spindle. Cell. 2000 Aug 18;102(4):399-402. Review. PubMed PMID: 10966101.

Merdes A, Heald R, Samejima K, Earnshaw WC, Cleveland DW. Formation of spindle poles by dynein/dynactin-dependent transport of NuMA. J Cell Biol. 2000 May 15;149(4):851-62. PubMed PMID: 10811826; PubMed Central PMCID: PMC2174573.

Heald R. A dynamic duo of microtubule modulators. Nat Cell Biol. 2000 Jan;2(1):E11-2. PubMed PMID: 10620815.

Heald R, Weis K. Spindles get the ran around. Trends Cell Biol. 2000 Jan;10(1):1-4. PubMed PMID: 10603469.


Rosania GR, Merlie J Jr, Gray N, Chang YT, Schultz PG, Heald R. A cyclin-dependent kinase inhibitor inducing cancer cell differentiation: biochemical identification using Xenopus egg extracts. Proc Natl Acad Sci U S A.  1999 Apr 27;96(9):4797-802. PubMed PMID: 10220373; PubMed Central PMCID: PMC21771.

Heald R, Walczak CE. Microtubule-based motor function in mitosis. Curr Opin Struct Biol. 1999 Apr;9(2):268-74. Review. PubMed PMID: 10322211.

Walczak CE, Vernos I, Mitchison TJ, Karsenti E, Heald R. A model for the proposed roles of different microtubule-based motor proteins in establishing spindle bipolarity. Curr Biol. 1998 Jul 30-Aug 13;8(16):903-13. PubMed PMID: 9707401.


Heald R, Tournebize R, Habermann A, Karsenti E, Hyman A. Spindle assembly in  Xenopus egg extracts: respective roles of centrosomes and microtubule self-organization. J Cell Biol. 1997 Aug 11;138(3):615-28. PubMed PMID: 9245790;  PubMed Central PMCID: PMC2141625.

Tournebize R, Heald R, Hyman A. Role of chromosomes in assembly of meiotic and mitotic spindles. Prog Cell Cycle Res. 1997;3:271-84. Review. PubMed PMID: 9552422.

Heald R, Tournebize R, Blank T, Sandaltzopoulos R, Becker P, Hyman A, Karsenti E. Self-organization of microtubules into bipolar spindles around artificial chromosomes in Xenopus egg extracts. Nature. 1996 Aug 1;382(6590):420-5. PubMed PMID: 8684481.

Heald R, McLoughlin M, McKeon F. Human wee1 maintains mitotic timing by protecting the nucleus from cytoplasmically activated Cdc2 kinase. Cell. 1993 Aug 13;74(3):463-74. PubMed PMID: 8348613.

Heald R, McKeon F. Mutations of phosphorylation sites in lamin A that prevent nuclear lamina disassembly in mitosis. Cell. 1990 May 18;61(4):579-89. PubMed PMID: 2344612.