Reitz, D., Djeghmoum, Y., Watson, R., Rajput, P., Argueso, L., Heyer, W.-D., and Piazza, A. (2023) Delineation of two multi-invasion-induced rearrangement pathways that differently affect genome stability. Genes Dev. 37, 621-639. PMID: 37541760.

Cerqueira P.G., Meyer D., Zhang L., Mallory B., Liu J., Hua Fu B.X., Zhang X., Heyer W.-D. (2023) Saccharomyces cerevisiae DNA polymerase IV overcomes Rad51 inhibition of DNA polymerase delta in Rad52-mediated direct-repeat recombination. Nucleic Acids Res 51:5547-5564. PMID: 37070185.


Reitz D., Savocco J., Piazza A., Heyer W.-D. (2022) Detection of Homologous Recombination Intermediates via Proximity Ligation and Quantitative PCR in Saccharomyces cerevisiae. J Vis Exp 187:10.3791/64240. PMID: 36155960.


Llorens-Agost M., Ensminger M., Le H.P., Heyer W.-D., Löbrich M. (2021) Turning end-joining upside down in mitosis. Mol Cell Oncol 8:2007029. PMID: 35419469.

Llorens-Agost M., Ensminger M., Le H.P., Gawai A., Liu J., Cruz-García A., Bhetawal S., Wood R.D., Heyer W.-D., Löbrich M. (2021) POLθ-mediated end joining is restricted by RAD52 and BRCA2 until the onset of mitosis. Nat Cell Biol 23, 1095–1104. PMID: 34616022.

Jay, A., Reitz, D., Namekawa, S.H., Heyer, W.-D. (2021) Cancer testis antigens and genomic instability: More than immunology. DNA Repair 108:103214. PMID: 34481156. 

Le, H.P., Heyer, W.-D., Liu, J. (2021) Guardians of the Genome: BRCA2 and Its Partners. Genes 12:1229. PMID: PMC8394001.


Shah S.S., Hartono S., Piazza A., Som V., Wright W., Chédin F., Heyer W.-D. (2020) Rdh54/Tid1 inhibits Rad51-Rad54-mediated D-loop formation and limits D-loop length. Elife 9:e59112. PMID: 33185188.

Shah S.S., Hartono S.R., Chédin F., Heyer W.-D. (2020) Bisulfite treatment and single-molecule real-time sequencing reveal D-loop length, position, and distribution. Elife 9:e59111. PMID: 33185185.

Schwartz E.K., Hung S.-H., Meyer D., Piazza A., Yan K., Fu B.X.H., Heyer W.-D. (2020) Mus81-Mms4 endonuclease is required for telomere maintenance under replication stress in Saccharomyces cerevisiae. PLoS Genetics 16:e1008816. PMID: 32469862.

Le H.P., Ma X., Vaquero J., Brinkmeyer M., Guo F., Heyer W.-D., and Liu J. (2020) DSS1 and ssDNA regulate oligomerization of BRCA2. Nucleic Acids Res 48:7818-7833. PMID: 32609828.

Piazza A., Rajput P., and Heyer W.-D. (2020) Physical and genetic assays for the study of DNA joint molecules metabolism and multi-invasion-induced rearrangements in S. cerevisiae. Methods Mol Biol 2153:535-554. PMID: 32840803.


Piazza A., Shah S.S., Wright W.D., Gore S.K., Koszul R., Heyer W.-D. (2019) Dynamic Processing of Displacement Loops during Recombinational DNA Repair. Mol Cell 73:1255-1266.e4. PMID: 30737186.

Meyer D., Fu B.X.H., Chavez M., Loeillet S., Cerqueira P.G., Nicolas A., Heyer W.-D. (2019) Cooperation between non-essential DNA polymerases contributes to genome stability in Saccharomyces cerevisiae. DNA Repair 76:40-49. PMID: 30818168.

Klein H.L., Ang K.K.H., Arkin M.R., Beckwitt E.C., Chang Y.-H., Fan J., Kwon Y., Morten M.J., Mukherjee S., Pambos O., el Sayyed H., Thrall E.S., Vieira-da-Rocha J., Wang Q., Wang S., Yeh H.-Y., Biteen Y.S., Chi P., Heyer W.-D., Kapanidis A., Loparo J.J., Strick T.R., Sung P., Van Houten B., Niu H., Rothenberg E. (2019) Guidelines for DNA recombination and repair studies: Mechanistic assay of DNA repair processes. Microb Cell 6:65-101. PMID: 30652106. 

Piazza A. and Heyer W.-D. (2019) Moving forward one step back at a time: reversibility during homologous recombination. Curr Genet 65:1333-1340. PMID: 31123771.

Jenkins, S.S., Gore, S., Guo, X., Liu, J., Ede, C., Veaute, X., Jinks-Robertson, S., Kowalczykowski, S.C., and Heyer, W.-D. (2019) Role of the Srs2-Rad51 interaction domain in crossover control in Saccharomyces cerevisiae. Genetics 212:1133-1145. PMID: 31142613.

Tavares, E.M., Wright, W.D., Heyer, W.-D., Le Cam, E., and Dupaigne, P. (2019) In vitro role of Rad54 in Rad51-ssDNA filament-dependent homology search and synaptic complexes formation. Nat Commun 10:4058. PMID: 31492866.


Piazza, A., and Heyer W.-D. (2018) Homologous Recombination and the Formation of Complex Genomic Rearrangements. Trends in Cell Biology. Volume 29, ISSUE 2, P135-149

Piazza, A., Koszul, R., and Heyer W.-D. (2018) A Proximity Ligation-based Method for Quantitative Measurement of D-loop Extension in S. cerevisiae. Meth. Enzymol. 601, 27-44.

Piazza, A. and Heyer,W.-D. (2018)Mechanisms of multi-invasion-induced Rearrangements: Implications for physiological and pathological recombination. BioEssays 40, e1700249.

Wright, W.D., Shah, S.S., and Heyer, W.-D. (2018) Homologous recombination and the repair of DNA double-strand breaks. J. Biol. Chem.Mar 29. pii: jbc.TM118.000372. doi: 10.1074/jbc.TM118.000372.


Piazza A, Wright WD, Heyer WD. Multi-invasions Are Recombination Byproducts that Induce Chromosomal Rearrangements. Cell. 2017 Aug 10;170(4):760-773.e15. doi: 10.1016 [PDF]

Liu J, Ede C, Wright WD, Gore SK, Jenkins SS, Freudenthal BD, Todd Washington M, Veaute X, Heyer WD. Srs2 promotes synthesis-dependent strand annealing by disrupting DNA polymerase δ-extending D-loops. Elife. 2017 May 23;6. pii: e22195. doi: 10.7554 (open access).


McVey M, Khodaverdian VY, Meyer D, Cerqueira PG, Heyer WD. Eukaryotic DNA Polymerases in Homologous Recombination. Annu Rev Genet. 2016 Nov 23;50:393-421. Review. [PDF]

Ganai RA, Zhang XP, Heyer WD, Johansson E. Strand displacement synthesis by yeast DNA polymerase ε. Nucleic Acids Res. 2016 Sep 30;44(17):8229-40. doi: 10.1093 (open access)

Spies J, Waizenegger A, Barton O, Sürder M, Wright WD, Heyer WD, Löbrich M. Nek1 Regulates Rad54 to Orchestrate Homologous Recombination and Replication Fork Stability. Mol Cell. 2016 Jun 1. pii: S1097-2765(16)30108-3. doi: 10.1016/j.molcel.2016.04.032. [PDF]

Janke, R., Braberg, H., Cantin, G., Yates III, J., Krogan, NJ, and Heyer, W.-D. (2016) Nonsense-mediated decay regulates key components of homologous recombination. Nucleic Acids Res.  doi: 10.1093/nar/gkw182 [Open Access]

Kowalczykowski, S.C., Hunter, N., and Heyer, W.-D. (2016) DNA Recombination. Perspectives in Biology, Cold Spring Harbor Press. [Buy]

Jenkins, S.S., Mukherjee, S., and Heyer, W.-D. (2016) DNA repair by homologous recombination. In: Ralph A Bradshaw and Philip D Stahl (Editors-in-Chief), Encyclopedia of Cell Biology, Vol 1, Waltham, MA: Academic Press, pp. 456-467. [PDF]

Crawley, J.N., Heyer, W.-D., and LaSalle, J.M. (2016) Autism and cancer share risk genes, pathways, and drug-targets. Trends in Genetics 32,139-146.[PDF]


Meyer D., Fu B., Heyer WD. DNA polymerases δ and λ cooperate in repairing double-strand breaks by microhomology-mediated end-joining in Saccharomyces cerevisiae, PNAS 2015, doi: 10.1073/pnas.1507833112 [PDF]

Heyer WD. Regulation of Recombination and Genomic Maintenance. Cold Spring Harb Perspect Biol. 2015 Aug 3;7(8). pii: a016501. doi: 10.1101/cshperspect.a016501 [PDF]

Mason JM, Dusad K, Wright WD, Grubb J, Budke B, Heyer WD, Connell PP, Weichselbaum RR, Bishop DK. RAD54 family translocases counter genotoxic effects of RAD51 in human tumor cells. Nucleic Acids Res. 2015 Mar 31;43(6):3180-96. doi: 10.1093/nar/gkv175. (open access)

Wright W.D. and Heyer W.-D. Recombination: DNA strand transferases. In Lennarz, W.J., and Lane, M.D. (eds.), Encyclopedia of Biological Chemistry, 2nd Edition, Elsevier (2013) [PDF]

Fasching CL, Cejka P, Kowalczykowski SC, Heyer WD Top3-rmi1 dissolve rad51-mediated d loops by a topoisomerase-based mechanism. Mol Cell. 2015 Feb 19;57(4):595-606. doi: 10.1016/j.molcel.2015.01.022. [PDF]


Mukherjee, S., Wright, W. D., Ehmsen, K. E., Heyer, W.-D. The Mus81-Mms4 structure-selective endonuclease requires nicked DNA junctions to undergo conformational changes and bends its DNA substrates for cleavage. Nucleic Acids Res, 2014 Apr 17. (open access)

Wright WD, Heyer WD. Rad54 functions as a Heteroduplex DNA Pump Modulated by Its DNA Substrates and Rad51 during D-loop Formation. Mol Cell. 2014 Feb 6;53(3):420-32. doi: 10.1016/j.molcel.2013.12.027. (free full text)


Zhang XP, Janke R, Kingsley J, Luo J, Fasching C, Ehmsen KT, Heyer WD. A Conserved Sequence Extending Motif III of the Motor Domain in the Snf2-Family DNA Translocase Rad54 Is Critical for ATPase Activity. PLoS One. 2013 Dec 16;8(12):e82184. doi: 10.1371/journal.pone.0082184 (open access)

Zinovyev A, Kuperstein I, Barillot E, Heyer WD. Synthetic lethality between gene defects affecting a single non-essential molecular pathway with reversible steps. PLoS Comput Biol. 2013 Apr;9(4):e1003016. (open access)

Sneeden JL, Grossi SM, Tappin I, Hurwitz J, Heyer WD. Reconstitution of recombination-associated DNA synthesis with human proteins. Nucleic Acids Res. 2013 May;41(9):4913-25. (open access)


Schwartz EK, Wright WD, Ehmsen KT, Evans JE, Stahlberg H, Heyer WD. Mus81-Mms4 functions as a single heterodimer to cleave nicked intermediates in recombinational DNA repair. Mol Cell Biol. 2012 Aug;32(15):3065-80. (free full text)

Muñoz-Galván S, Tous C, Blanco MG, Schwartz EK, Ehmsen KT, West SC, Heyer WD, Aguilera A. Distinct roles of Mus81, Yen1, Slx1-Slx4, and Rad1 nucleases in the repair of replication-born double-strand breaks by sister chromatid exchange. Mol Cell Biol. 2012 May;32(9):1592-603. (free full text)


Liu J, Renault L, Veaute X, Fabre F, Stahlberg H, Heyer WD. Rad51 paralogues Rad55-Rad57 balance the antirecombinase Srs2 in Rad51 filament formation. Nature. 2011 Oct 23;479(7372):245-8. (PDF)

Ceballos SJ, Heyer WD. Functions of the Snf2/Swi2 family Rad54 motor protein in homologous recombination. Biochim Biophys Acta. 2011 Sep;1809(9):509-23.

Liu J, Sneeden J, Heyer WD. In vitro assays for DNA pairing and recombination-associated DNA synthesis. Methods Mol Biol. 2011;745:363-83.

Wright WD, Ehmsen KT, Heyer WD. Assays for structure-selective DNA endonucleases. Methods Mol Biol. 2011;745:345-62.

Zhang XP, Heyer WD. Quality control of purified proteins involved in homologous recombination. Methods Mol Biol. 2011;745:329-43.

Liu J, Ehmsen KT, Heyer WD, Morrical SW. Presynaptic filament dynamics in homologous recombination and DNA repair. Crit Rev Biochem Mol Biol. 2011 Jun;46(3):240-70.

Schwartz EK, Heyer WD. Processing of joint molecule intermediates by structure-selective endonucleases during homologous recombination in eukaryotes. Chromosoma. 2011 Apr;120(2):109-27.

Liu J, Heyer WD. Who’s who in human recombination: BRCA2 and RAD52. PNAS. 2011 Jan 11;108(2):441-2.


Liu J, Doty T, Gibson B, Heyer WD. Human BRCA2 protein promotes RAD51 filament formation on RPA-covered single-stranded DNA. Nat Struct Mol Biol. 2010 Oct;17(10):1260-2. (PDF)

Heyer WD, Ehmsen KT, Liu J. Regulation of homologous recombination in eukaryotes. Annu Rev Genet. 2010;44:113-39. (PDF)

Janke R, Herzberg K, Rolfsmeier M, Mar J, Bashkirov VI, Haghnazari E, Cantin G, Yates JR 3rd, Heyer WD. A truncated DNA-damage-signaling response is activated after DSB formation in the G1 phase of Saccharomyces cerevisiae. Nucleic Acids Res. 2010 Apr;38(7):2302-13.


Li X, Stith CM, Burgers PM, Heyer WD. PCNA is required for initiation of recombination-associated DNA synthesis by DNA polymerase delta. Mol Cell. 2009 Nov 25;36(4):704-13.

Ehmsen KT, Heyer WD. A junction branch point adjacent to a DNA backbone nick directs substrate cleavage by Saccharomyces cerevisiae Mus81-Mms4. Nucleic Acids Res. 2009 Apr;37(6):2026-36.

Li X, Heyer WD. RAD54 controls access to the invading 3′-OH end after RAD51-mediated DNA strand invasion in homologous recombination in Saccharomyces cerevisiae. Nucleic Acids Res. 2009 Feb;37(2):638-46.

Zhang XP, Galkin VE, Yu X, Egelman EH, Heyer WD. Loop 2 in Saccharomyces cerevisiae Rad51 protein regulates filament formation and ATPase activity. Nucleic Acids Res. 2009 Jan;37(1):158-71.


Ehmsen KT, Heyer WD. Biochemistry of Meiotic Recombination: Formation, Processing, and Resolution of Recombination Intermediates. Genome Dyn Stab. 2008 Apr 5;3:91.

Ehmsen, K. and Heyer, WD. Saccharomyces cerevisiae Mus81-Mms4 is a catalytic, DNA structure-selective endonuclease Nucleic Acids Res. 36:2182.
This paper has been selected for Faculty of 1000 Biology. See the Evaluations.

Li, X. and Heyer, WD. Homologous recombination in DNA repair and DNA damage tolerance. Cell Res. 2008 Jan;18(1):99-113. doi: 10.1038/cr.2008.1.


Heyer, WD. Biochemistry of eukaryotic homologous recombination. In Topics in Current Genetics. vol 17: Molecular Genetics of Recombination 2007 Mar 1;17:95-133. (PDF)

Li, X., Zhang, X.-P., Solinger, J., Kiianitsa, K., Yu, X., Egelman, E. and Heyer, WD. Rad51 and Rad54 ATPase activites are both required to modulate Rad51-dsDNA filament dynamics.
Nucleic Acid Res. 2007;35(12):4124-40. Epub 2007 Jun 12.

Mimida N, Kitamoto H, Osakabe K, Nakashima M, Ito Y, Heyer WD., Toki S, Ichikawa H. Two alternatively spliced transcripts generated from OsMUS81, a rice homolog of yeast MUS81, are up-regulated by DNA-damaging treatments. Plant Cell Physiol. 2007 Apr;48(4):648-54. Epub 2007 Feb 27.


Symington LS, Heyer WD. Some disassembly required: role of DNA translocases in the disruption of recombination intermediates and dead-end complexes. Genes Dev. 2006 Sep 15;20(18):2479-86

Herzberg K, Bashkirov VI, Rolfsmeier M, Haghnazari E, McDonald WH, Anderson S, Bashkirova EV, Yates JR 3rd, Heyer WD. Phosphorylation of Rad55 on serines 2, 8, and 14 is required for efficient homologous recombination in the recovery of stalled replication forks. Mol Cell Biol. 2006 Nov;26(22):8396-409. Epub 2006 Sep 11.

Heyer WD, Li X, Rolfsmeier M, Zhang XP. Rad54: the Swiss Army knife of homologous recombination? Nucleic Acids Res. 2006;34(15):4115-25. Epub 2006 Aug 25

Bashkirov VI, Herzberg K, Haghnazari E, Vlasenko AS, Heyer WD. DNA damage-induced phosphorylation of Rad55 protein as a sentinel for DNA damage checkpoint activation in S. cerevisiae. Methods Enzymol. 2006;409:166-82.

Kiianitsa K, Solinger JA, Heyer WD. Terminal association of Rad54 protein with the Rad51-dsDNA filament. PNAS. 2006 Jun 27;103(26):9767-72. Epub 2006 Jun 19.

Galkin VE, Wu Y, Zhang XP, Qian X, He Y, Yu X, Heyer WD, Luo Y, Egelman EH. The Rad51/RadA N-terminal domain activates nucleoprotein filament ATPase activity. Structure. 2006 Jun;14(6):983-92.

Chin JK, Bashkirov VI, Heyer WD, Romesberg FE. Esc4/Rtt107 and the control of recombination during replication. DNA Repair (Amst). 2006 May 10;5(5):618-28. Epub 2006 Mar 29.


Zhang XP, Lee KI, Solinger JA, Kiianitsa K, Heyer WD. Gly-103 in the N-terminal domain of Saccharomyces cerevisiae Rad51 protein is critical for DNA binding. J Biol Chem. 2005 Jul 15;280(28):26303-11. Epub 2005 May 21.


Heyer WD. Damage signaling: RecQ sends an SOS to you. Curr Biol. 2004 Oct 26;14(20):R895-7.

Heyer WD, Kanaar R. Recombination mechanisms; fortieth anniversary meeting of the Holliday model. Mol Cell. 2004 Oct 8;16(1):1-9.

Haghnazari E, Heyer WD. The DNA damage checkpoint pathways exert multiple controls on the efficiency and outcome of the repair of a double-stranded DNA gap. Nucleic Acids Res. 2004 Aug 10;32(14):4257-68.

Haghnazari E, Heyer WD. The Hog1 MAP kinase pathway and the Mec1 DNA damage checkpoint pathway independently control the cellular responses to hydrogen peroxide. DNA Repair (Amst). 2004 Jul 2;3(7):769-76.

Heyer WD. A new deal for Holliday junctions. Nat Struct Mol Biol. 2004 Feb;11(2):117-9.

Heyer WD. Recombination: Holliday junction resolution and crossover formation. Curr Biol. 2004 Jan 20;14(2):R56-8


Heyer WD, Ehmsen KT, Solinger JA. Holliday junctions in the eukaryotic nucleus: resolution in sight? Trends Biochem Sci. 2003 Oct;28(10):548-57

Kiianitsa K, Solinger JA, Heyer WD. NADH-coupled microplate photometric assay for kinetic studies of ATP-hydrolyzing enzymes with low and high specific activities. Anal Biochem. 2003 Oct 15;321(2):266-71

Mallory JC, Bashkirov VI, Trujillo KM, Solinger JA, Dominska M, Sung P, Heyer WD, Petes TD. Amino acid changes in Xrs2p, Dun1p, and Rfa2p that remove the preferred targets of the ATM family of protein kinases do not affect DNA repair or telomere length in Saccharomyces cerevisiae. DNA Repair (Amst). 2003 Sep 18;2(9):1041-64.

Schmuckli-Maurer J, Rolfsmeier M, Nguyen H, Heyer WD. Genome instability in rad54 mutants of Saccharomyces cerevisiae. Nucleic Acids Res. 2003 Feb 1;31(3):1013-23.

Bashkirov VI, Bashkirova EV, Haghnazari E, Heyer WD. Direct kinase-to-kinase signaling mediated by the FHA phosphoprotein recognition domain of the Dun1 DNA damage checkpoint kinase. Mol Cell Biol. 2003 Feb;23(4):1441-52.


Fabre F, Chan A, Heyer WD, Gangloff S. Alternate pathways involving Sgs1/Top3, Mus81/ Mms4, and Srs2 prevent formation of toxic recombination intermediates from single-stranded gaps created by DNA replication. Proc Natl Acad Sci U S A. 2002 Dec 24;99(26):16887-92. Epub 2002 Dec 10. Erratum in: Proc Natl Acad Sci U S A. 2003 Feb 4;100(3):1462.

Solinger JA, Kiianitsa K, Heyer WD. Rad54, a Swi2/Snf2-like recombinational repair protein, disassembles Rad51:dsDNA filaments. Mol Cell. 2002 Nov;10(5):1175-88.

Kiianitsa K, Solinger JA, Heyer WD. Rad54 protein exerts diverse modes of ATPase activity on duplex DNA partially and fully covered with Rad51 protein. J Biol Chem. 2002 Nov 29;277(48):46205-15. Epub 2002 Sep 30.

Walsh L, Schmuckli-Maurer J, Billinton N, Barker MG, Heyer WD, Walmsley RM. DNA-damage induction of RAD54 can be regulated independently of the RAD9- and DDC1-dependent checkpoints that regulate RNR2. Curr Genet. 2002 Jul;41(4):232-40. Epub 2002 Jun 27.

Kim PM, Paffett KS, Solinger JA, Heyer WD, Nickoloff JA. Spontaneous and double-strand break-induced recombination, and gene conversion tract lengths, are differentially affected by overexpression of wild-type or ATPase-defective yeast Rad54. Nucleic Acids Res. 2002 Jul 1;30(13):2727-35.


Haber JE, Heyer WD. The fuss about Mus81.Cell. 2001 Nov 30;107(5):551-4.

Solinger JA, Heyer WD. Rad54 protein stimulates the postsynaptic phase of Rad51 protein-mediated DNA strand exchange. Proc Natl Acad Sci U S A. 2001 Jul 17;98(15):8447-53.

Solinger JA, Lutz G, Sugiyama T, Kowalczykowski SC, Heyer WD. Rad54 protein stimulates heteroduplex DNA formation in the synaptic phase of DNA strand exchange via specific interactions with the presynaptic Rad51 nucleoprotein filament. J Mol Biol. 2001 Apr 13;307(5):1207-21.


Boddy MN, Lopez-Girona A, Shanahan P, Interthal H, Heyer WD, Russell P. Damage tolerance protein Mus81 associates with the FHA1 domain of checkpoint kinase Cds1. Mol Cell Biol. 2000 Dec;20(23):8758-66.

Mazin AV, Bornarth CJ, Solinger JA, Heyer WD, Kowalczykowski SC. Rad54 protein is targeted to pairing loci by the Rad51 nucleoprotein filament. Mol Cell. 2000 Sep;6(3):583-92.

Gos P, Eicher B, Kohli J, Heyer WD. No mutagenic or recombinogenic effects of mobile phone fields at 900 MHz detected in the yeast Saccharomyces cerevisiae. Bioelectromagnetics. 2000 Oct;21(7):515-23.

Interthal H, Heyer WD. MUS81 encodes a novel helix-hairpin-helix protein involved in the response to UV- and methylation-induced DNA damage in Saccharomyces cerevisiae. Mol Gen Genet. 2000 Jun;263(5):812-27.

Schmuckli-Maurer J, Heyer WD. Meiotic recombination in RAD54 mutants of Saccharomyces cerevisiae. Chromosoma. 2000;109(1-2):86-93.

Bashkirov VI, King JS, Bashkirova EV, Schmuckli-Maurer J, Heyer WD. DNA repair protein Rad55 is a terminal substrate of the DNA damage checkpoints. Mol Cell Biol. 2000 Jun;20(12):4393-404.


Solinger JA, Pascolini D, Heyer WD. Active-site mutations in the Xrn1p exoribonuclease of Saccharomyces cerevisiae reveal a specific role in meiosis. Mol Cell Biol. 1999 Sep;19(9):5930-42.

Khasanov FK, Savchenko GV, Bashkirova EV, Korolev VG, Heyer WD, Bashkirov VI. A new recombinational DNA repair gene from Schizosaccharomyces pombe with homology to Escherichia coli RecA. Genetics. 1999 Aug;152(4):1557-72.

Clever B, Schmuckli-Maurer J, Sigrist M, Glassner BJ, Heyer WD. Specific negative effects resulting from elevated levels of the recombinational repair protein Rad54p in Saccharomyces cerevisiae. Yeast. 1999 Jun 30;15(9):721-40.

Schmuckli-Maurer J, Heyer WD. The Saccharomyces cerevisiae RAD54 gene is important but not essential for natural homothallic mating-type switching. Mol Gen Genet. 1999 Jan;260(6):551-8.


Billinton N, Barker MG, Michel CE, Knight AW, Heyer WD, Goddard NJ, Fielden PR, Walmsley RM. Development of a green fluorescent protein reporter for a yeast genotoxicity biosensor. Biosens Bioelectron. 1998 Oct 1;13(7-8):831-8.


Walmsley RM, Billinton N, Heyer WD. Green fluorescent protein as a reporter for the DNA damage-induced gene RAD54 in Saccharomyces cerevisiae. Yeast. 1997 Dec;13(16):1535-45.

Clever B, Interthal H, Schmuckli-Maurer J, King J, Sigrist M, Heyer WD. Recombinational repair in yeast: functional interactions between Rad51 and Rad54 proteins. EMBO J. 1997 May 1;16(9):2535-44.

Bashkirov VI, Scherthan H, Solinger JA, Buerstedde JM, Heyer WD. A mouse cytoplasmic exoribonuclease (mXRN1p) with preference for G4 tetraplex substrates. J Cell Biol. 1997 Feb 24;136(4):761-73.

Gos P, Eicher B, Kohli J, Heyer WD. Extremely high frequency electromagnetic fields at low power density do not affect the division of exponential phase Saccharomyces cerevisiae cells. Bioelectromagnetics. 1997;18(2):142-55.


Kanaar R, Troelstra C, Swagemakers SM, Essers J, Smit B, Franssen JH, Pastink A, Bezzubova OY, Buerstedde JM, Clever B, Heyer WD, Hoeijmakers JH. Human and mouse homologs of the Saccharomyces cerevisiae RAD54 DNA repair gene: evidence for functional conservation. Curr Biol. 1996 Jul 1;6(7):828-38.

Santos-Rosa H, Clever B, Heyer WD, Aguilera A. The yeast HRS1 gene encodes a polyglutamine-rich nuclear protein required for spontaneous and hpr1-induced deletions between direct repeats. Genetics. 1996 Mar;142(3):705-16


Bashkirov VI, Solinger JA, Heyer WD. Identification of functional domains in the Sep1 protein (= Kem1, Xrn1), which is required for transition through meiotic prophase in Saccharomyces cerevisiae. Chromosoma. 1995 Nov;104(3):215-22.

Holler A, Bashkirov VI, Solinger JA, Reinhart U, Heyer WD. Use of monoclonal antibodies in the functional characterization of the Saccharomyces cerevisiae Sep1 protein. Eur J Biochem. 1995 Jul 15;231(2):329-36.

Heyer WD, Johnson AW, Reinhart U, Kolodner RD. Regulation and intracellular localization of Saccharomyces cerevisiae strand exchange protein 1 (Sep1/Xrn1/Kem1), a multifunctional exonuclease. Mol Cell Biol. 1995 May;15(5):2728-36.

Interthal H, Bellocq C, Bähler J, Bashkirov VI, Edelstein S, Heyer WD. A role of Sep1 (= Kem1, Xrn1) as a microtubule-associated protein in Saccharomyces cerevisiae. EMBO J. 1995 Mar 15;14(6):1057-66.


Käslan E, Heyer WD. Schizosaccharomyces pombe fatty acid synthase mediates DNA strand exchange in vitro. J Biol Chem. 1994 May 13;269(19):14103-10.

Käslin E, Heyer WD. A multifunctional exonuclease from vegetative Schizosaccharomyces pombe cells exhibiting in vitro strand exchange activity. J Biol Chem. 1994 May 13;269(19):14094-102.

Bähler J, Hagens G, Holzinger G, Scherthan H, Heyer WD. Saccharomyces cerevisiae cells lacking the homologous pairing protein p175SEP1 arrest at pachytene during meiotic prophase. Chromosoma. 1994 Apr;103(2):129-41.

Heyer WD. The search for the right partner: homologous pairing and DNA strand exchange proteins in eukaryotes. Experientia. 1994 Mar 15;50(3):223-33.

Heyer WD, Kohli J. Homologous recombination. Experientia. 1994 Mar 15;50(3):189-91.


Bezzubova OY, Schmidt H, Ostermann K, Heyer WD, Buerstedde JM. Identification of a chicken RAD52 homologue suggests conservation of the RAD52 recombination pathway throughout the evolution of higher eukaryotes. Nucleic Acids Res. 1993 Dec 25;21(25):5945-9.

Post-doc years:

Heyer WD, Kolodner RD. Enzymology of homologous recombination in Saccharomyces cerevisiae. Prog Nucleic Acid Res Mol Biol. 1993;46:221-71.

Erdile LF, Heyer WD, Kolodner R, Kelly TJ. Characterization of a cDNA encoding the 70-kDa single-stranded DNA-binding subunit of human replication protein A and the role of the protein in DNA replication. J Biol Chem. 1991 Jun 25;266(18):12090-8.

Heyer WD, Johnson AW, Norris DN, Tishkoff D, Kolodner RD. Saccharomyces cerevisiae proteins involved in hybrid DNA formation in vitro. Biochimie. 1991 Feb-Mar;73(2-3):269-76.

Heyer WD, Rao MR, Erdile LF, Kelly TJ, Kolodner RD. An essential Saccharomyces cerevisiae single-stranded DNA binding protein is homologous to the large subunit of human RP-A. EMBO J. 1990 Jul;9(7):2321-9.

Heyer WD, Kolodner RD. Purification and characterization of a protein from Saccharomyces cerevisiae that binds tightly to single-stranded DNA and stimulates a cognate strand exchange protein. Biochemistry. 1989 Apr 4;28(7):2856-62.

Szankasi P, Heyer WD, Schuchert P, Kohli J. DNA sequence analysis of the ade6 gene of Schizosaccharomyces pombe. Wild-type and mutant alleles including the recombination host spot allele ade6-M26. J Mol Biol. 1988 Dec 20;204(4):917-25.

Heyer WD, Evans DH, Kolodner RD. Renaturation of DNA by a Saccharomyces cerevisiae protein that catalyzes homologous pairing and strand exchange. J Biol Chem. 1988 Oct 15;263(29):15189-95.

Graduate years:

Grossenbacher AM, Stadelmann B, Heyer WD, Thuriaux P, Kohli J, Smith C, Agris PF, Kuo KC, Gehrke C. Antisuppressor mutations and sulfur-carrying nucleosides in transfer RNAs of Schizosaccharomyces pombe. J Biol Chem. 1986 Dec 15;261(35):16351-5.

Willis I, Nichols M, Chisholm V, Söll D, Heyer WD, Szankasi P, Amstutz H, Munz P, Kohli J. Functional complementation between mutations in a yeast suppressor tRNA gene reveals potential for evolution of tRNA sequences. PNAS. 1986 Oct;83(20):7860-4.

Heyer WD, Munz P, Amstutz H, Aebi R, Gysler C, Schuchert P, Szankasi P, Leupold U, Kohli J, Gamulin V, and Söll, D. Inactivation of nonsense suppressor transfer RNA genes in Schizosaccharomyces pombe. Intergenic conversion and hot spots of mutation. J Mol Biol. 1986 Apr 5;188(3):343-53.

Wright A, Maundrell K, Heyer WD, Beach D, Nurse P. Vectors for the construction of gene banks and the integration of cloned genes in Schizosaccharomyces pombe and Saccharomyces cerevisiae. Plasmid. 1986 Mar;15(2):156-8.

Heyer WD, Sipiczki M, Kohli J. Replicating plasmids in Schizosaccharomyces pombe: improvement of symmetric segregation by a new genetic element. Mol Cell Biol. 1986 Jan;6(1):80-9

Amstutz H, Munz P, Heyer WD, Leupoid U, Kohli J. Concerted evolution of tRNA genes: intergenic conversion among three unlinked serine tRNA genes in S. pombe. Cell. 1985 Apr;40(4):879-86.

Heyer WD, Thuriaux P, Kohli J, Ebert P, Kersten H, Gehrke C, Kuo KC, Agris PF. An antisuppressor mutation of Schizosaccharomyces pombe affects the post-transcriptional modification of the “wobble” base in the anticodon of tRNAs. J Biol Chem. 1984 Mar 10;259(5):2856-62.

Kohli J, Munz P, Aebi R, Amstutz H, Gysler C, Heyer WD, Lehmann L, Schuchert P, Szankasi P, Thuriaux P, Leupold U, Bell J, Gamulin V, Hottinger H, Pearson D and Söll D. Interallelic and intergenic conversion in three serine tRNA genes of Schizosaccharomyces pombe. Cold Spring Harb Symp Quant Biol. 1984;49:31-40.

Thuriaux P, Heyer WD, Strauss A. Organisation of the complex locus trp1 in the fission yeast Schizosaccharomyces pombe. Curr Genet. 1982 Oct;6(1):13-8. doi: 10.1007/BF00397634.


Walmsley, R. M. and Heyer, W.-D. (1998) Detection of DNA damage ‘GFP’.
UK Patent 9706414.1