Microbiology and Molecular Genetics | UC Davis


Yun Y, Lee S, So C, Manhas R, Kim C, Wibowo T, Hori M, Hunter N. Oocyte Development and Quality in Young and Old Mice following Exposure to Atrazine. Environ Health Perspect. 2022 Nov;130(11):117007. doi: 10.1289/EHP11343. Epub 2022 Nov 11. PMID: 36367780; PMCID: PMC9651182.


Sandhu S., Salmon L.J., Hunter J.E. Wilson C.L., Perkins N.D., Hunter N., Davies O.R., McClurg U.L.(2021). Physiological and pathological roles for synaptonemal complex protein TEX12 in meiotic and mitotic microtubule organizing centers. In revision at Communications Biology.
† Sumit Sandhu from the Hunter lab is joint first author.

He, W., Verhees, G., Bhagwat, N., Yang, Y., Kulkarni, D., Lombardo, Z., Lahiri, S., Roy, P., Zhuo, J., Dang, B., Snyder, A., Shastry, S., Moezpoor, M., Alocozy, L., Lee, K.G., Painter, D., Mukerji, I., Hunter, N. (2021) SUMO Fosters Assembly of the MutSƔ Complex to Facilitate Meiotic Crossing Over. Developmental Cell. doi: 10.1016/j.devcel.2021.06.012. PMID: 34214491

Bozdag, G.O., Ono, J., Denton, J.A., Karakoc, E., Hunter, N., Leu, J-Y. and Greig, D. (2021). Breaking a species barrier by enabling hybrid recombination. Current Biology 31(4):R180-R181.

Bhagwat, N. R., Owens, S. N., Ito, M., Boinapalli, J. V., Poa, P., Ditzel, A., Kopparapu, S., Mahalawat, M., Davies, O. R., Collins, S. R., Johnson, J. R., Krogan, N. J., & Hunter, N. (2021). SUMO is a pervasive regulator of meiosis. eLife10, e57720.


Yun, Y., Ito, M., Sandhu, S. and and Hunter, N. (2020). Cytological monitoring of meiotic crossovers in spermatocytes and oocytes. Homologous Recombination: Methods and Protocols, Editors Andres Aguilera, PhD and Aura Carreira. Methods in Molecular Biology. 2153, 267-286.

Kulkarni D, Owens S, Honda M, Ito, M, Yang Y, Corrigan M, Chen L, Quan A, Hunter N. PCNA activates the MutLɣ endonuclease to promote meiotic crossing over. Nature, 586, 623-627.

Bondarieva, A., Raveendran, K., Rao, H.B.D.P., Ravindranathan, R., Zorzompokou, C., Finsterbusch, F., Dereli, I., Telychko, V., Tränkner, D., Schleiffer, A., Fei, J-F., Klimova, A., Ito, M., Kulkarni, D.S. Roeder, I., Hunter, N., Tóth, A. (2020). Proline-rich protein PRR19 functions with cyclin-like CNTD1 to promote meiotic crossing over in mouse. Nature Communications 18, 3101.

He, W., Rao, H., Tang, S., Bhagwat, N., Kulkarni, D. S., Ma, Y., Chang, M., Hall, C., Bragg, J. W., Manasca, H. S., Baker, C., Verhees, G. F., Ranjha, L., Chen, X., Hollingsworth, N. M., Cejka, P., & Hunter, N. (2020). Regulated Proteolysis of MutSγ Controls Meiotic Crossing Over. Molecular cell78(1), 168–183.e5.


Yun, Y., Wei, Z., & Hunter, N. (2019). Maternal obesity enhances oocyte chromosome abnormalities associated with aging. Chromosoma128(3), 413–421.


Qiao, H., Rao, H., Yun, Y., Sandhu, S., Fong, J. H., Sapre, M., Nguyen, M., Tham, A., Van, B. W., Chng, T., Lee, A., & Hunter, N. (2018). Impeding DNA Break Repair Enables Oocyte Quality Control. Molecular cell72(2), 211–221.e3.

Owens, S., Tang, S., & Hunter, N. (2018). Monitoring Recombination During Meiosis in Budding Yeast. Methods in enzymology601, 275–307.


Rao, H. B., Qiao, H., Bhatt, S. K., Bailey, L. R., Tran, H. D., Bourne, S. L., Qiu, W., Deshpande, A., Sharma, A. N., Beebout, C. J., Pezza, R. J., & Hunter, N. (2017). A SUMO-ubiquitin relay recruits proteasomes to chromosome axes to regulate meiotic recombination. Science (New York, N.Y.)355(6323), 403–407.

Hunter N. (2017). Oocyte Quality Control: Causes, Mechanisms, and Consequences. Cold Spring Harbor symposia on quantitative biology82, 235–247.


Hunter N. (2015). Meiotic Recombination: The Essence of Heredity. Cold Spring Harbor perspectives in biology7(12), a016618.

DNA Recombination. Cold Spring Harbor Perspectives in Biology. (2015). Edited by Stephen Kowalczykowski, Neil Hunter and Wolf-Dietrich Heyer. Cold Spring Harbor Press. Buy the book!

Tang, S., Wu, M., Zhang, R., & Hunter, N. (2015). Pervasive and essential roles of the Top3-Rmi1 decatenase orchestrate recombination and facilitate chromosome segregation in meiosis. Molecular cell57(4), 607–621.


Qiao, H., Prasada Rao, H. B., Yang, Y., Fong, J. H., Cloutier, J. M., Deacon, D. C., Nagel, K. E., Swartz, R. K., Strong, E., Holloway, J. K., Cohen, P. E., Schimenti, J., Ward, J., & Hunter, N. (2014). Antagonistic roles of ubiquitin ligase HEI10 and SUMO ligase RNF212 regulate meiotic recombination. Nature genetics46(2), 194–199.


Copsey, A., Tang, S., Jordan, P. W., Blitzblau, H. G., Newcombe, S., Chan, A. C., Newnham, L., Li, Z., Gray, S., Herbert, A. D., Arumugam, P., Hochwagen, A., Hunter, N., & Hoffmann, E. (2013). Smc5/6 coordinates formation and resolution of joint molecules with chromosome morphology to ensure meiotic divisions. PLoS genetics9(12), e1004071.

Lao, J. P., Cloud, V., Huang, C. C., Grubb, J., Thacker, D., Lee, C. Y., Dresser, M. E., Hunter, N., & Bishop, D. K. (2013). Meiotic crossover control by concerted action of Rad51-Dmc1 in homolog template bias and robust homeostatic regulation. PLoS genetics9(12), e1003978.

Sasanuma, H., Tawaramoto, M. S., Lao, J. P., Hosaka, H., Sanda, E., Suzuki, M., Yamashita, E., Hunter, N., Shinohara, M., Nakagawa, A., & Shinohara, A. (2013). A new protein complex promoting the assembly of Rad51 filaments. Nature communications4, 1676.

Lao, J.P., Tang, S. and Hunter, N. (2013). Native/Denaturing Two-Dimensional DNA Electrophoresis And Its Application To The Analysis Of Recombination Intermediates. In, Methods in Molecular Biology, DNA Electrophoresis: Methods and Protocols (Totowa, Humana Press).

Reynolds, A., Qiao, H., Yang, Y., Chen, J. K., Jackson, N., Biswas, K., Holloway, J. K., Baudat, F., de Massy, B., Wang, J., Höög, C., Cohen, P. E., & Hunter, N. (2013). RNF212 is a dosage-sensitive regulator of crossing-over during mammalian meiosis. Nature genetics45(3), 269–278.


Biswas, K., Das, R., Eggington, J.M., Qiao, H., North, S.L., Stauffer, S., Burkett, S., Betty K. Martin, B.S., Southon, E, Sizemore, S.C., Pruss, D., Bowles, K.R., Roa, B.B., Hunter, N., Tessarollo, L., Wenstrup, R.J., Byrd, R.A. and Sharan, S.K. (2012). Functional evaluation of BRCA2 variants mapping to the PALB2 binding and C-terminal DNA binding domains using a mouse ES cell–based assay. Human Molecular Genetics 21, 3993-4006. 

Qiao H., Chen, J., Reynolds, A., Höög, C., Paddy, M. and Hunter, N. (2012). Interplay between centromeres, synaptonemal complex and homologous recombination during mammalian meiosis. PLoS Genetics, PLoS Genet 8(6): e1002790.

Zakheryevich, K, Tang, S., Ma, Y. and Hunter, N. (2012). Delineation of joint molecule resolution pathways in meiosis identifies a crossover-specific resolvase. Cell. 2012;149(2):334-347.


Hunter N. (2011). Double duty for Exo1 during meiotic recombination. Cell cycle (Georgetown, Tex.)10(16), 2607–2609.

Hwang, P. Y., & Hunter, N. (2011). Mapping meiotic breaks: Spo11 oligonucleotides precisely mark the spots. Genome biology12(4), 111.


Zakharyevich, K., Ma, Y., Tang, S., Hwang, P. Y., Boiteux, S., & Hunter, N. (2010). Temporally and biochemically distinct activities of Exo1 during meiosis: double-strand break resection and resolution of double Holliday junctions. Molecular cell40(6), 1001–1015.

Hunter, N. (2010). Meiosis. The Encyclopedia of Biological Chemistry 2nd Edition, Lennarz, W. and Lane, M. (Eds), Elsevier Press, San Diego.

Lao, J. P., & Hunter, N. (2010). Trying to avoid your sister. PLoS biology8(10), e1000519.

Bzymek, M., Thayer, N. H., Oh, S. D., Kleckner, N., & Hunter, N. (2010). Double Holliday junctions are intermediates of DNA break repair. Nature464(7290), 937–941.


Oh, S. D., Jessop, L., Lao, J. P., Allers, T., Lichten, M., & Hunter, N. (2009). Stabilization and electrophoretic analysis of meiotic recombination intermediates in Saccharomyces cerevisiae. Methods in molecular biology (Clifton, N.J.)557, 209–234.


Oh, S. D., Lao, J. P., Taylor, A. F., Smith, G. R., & Hunter, N. (2008). RecQ helicase, Sgs1, and XPF family endonuclease, Mus81-Mms4, resolve aberrant joint molecules during meiotic recombination. Molecular cell31(3), 324–336.

Hunter N. (2008). Hop1 and the meiotic DNA-damage response. Cell132(5), 731–732.

Hunter N. (2008). The RecQ DNA helicases: Jacks-of-all-trades or master-tradesmen?. Cell research18(3), 328–330.

Shinohara, M., Oh, S. D., Hunter, N., & Shinohara, A. (2008). Crossover assurance and crossover interference are distinctly regulated by the ZMM proteins during yeast meiosis. Nature genetics40(3), 299–309.

Lao, J. P., Oh, S. D., Shinohara, M., Shinohara, A., & Hunter, N. (2008). Rad52 promotes postinvasion steps of meiotic double-strand-break repair. Molecular cell29(4), 517–524.


Oh, S. D., Lao, J. P., Hwang, P. Y., Taylor, A. F., Smith, G. R., & Hunter, N. (2007). BLM ortholog, Sgs1, prevents aberrant crossing-over by suppressing formation of multichromatid joint molecules. Cell130(2), 259–272.


Hunter, N. (2006). Meiotic Recombination. Topics in Current Genetics, Molecular Genetics of Recombination, Aguilera, A. and Rothstein, R. (Eds), Springer-Verlag, Heidelberg, pp. 381-442.

Cromie, G. A., Hyppa, R. W., Taylor, A. F., Zakharyevich, K., Hunter, N., & Smith, G. R. (2006). Single Holliday junctions are intermediates of meiotic recombination. Cell127(6), 1167–1178.

Martini, E., Diaz, R. L., Hunter, N., & Keeney, S. (2006). Crossover homeostasis in yeast meiosis. Cell126(2), 285–295.


McDougall, A., Elliott, D. J., & Hunter, N. (2005). Pairing, connecting, exchanging, pausing and pulling chromosomes. EMBO reports6(2), 120–125.


Hunter, N. (2004). Meiosis. The Encyclopedia of Biological Chemistry, 1st Edition, Lennarz, W. and Lane, M. (Eds), Elsevier Press, San Diego, pp. 610-616. 

Börner, G. V., Kleckner, N., & Hunter, N. (2004). Crossover/noncrossover differentiation, synaptonemal complex formation, and regulatory surveillance at the leptotene/zygotene transition of meiosis. Cell117(1), 29–45.


Hunter N. (2003). Synaptonemal complexities and commonalities. Molecular cell12(3), 533–535.

de los Santos, T., Hunter, N., Lee, C., Larkin, B., Loidl, J., & Hollingsworth, N. M. (2003). The Mus81/Mms4 endonuclease acts independently of double-Holliday junction resolution to promote a distinct subset of crossovers during meiosis in budding yeast. Genetics164(1), 81–94.


Blat, Y., Protacio, R. U., Hunter, N., & Kleckner, N. (2002). Physical and functional interactions among basic chromosome organizational features govern early steps of meiotic chiasma formation. Cell111(6), 791–802.


Hunter, N., & Kleckner, N. (2001). The single-end invasion: an asymmetric intermediate at the double-strand break to double-holliday junction transition of meiotic recombination. Cell106(1), 59–70.

Hunter, N., Börner, G. V., Lichten, M., & Kleckner, N. (2001). Gamma-H2AX illuminates meiosis. Nature genetics27(3), 236–238.

1999 and earlier

Wang, T. F., Kleckner, N., & Hunter, N. (1999). Functional specificity of MutL homologs in yeast: evidence for three Mlh1-based heterocomplexes with distinct roles during meiosis in recombination and mismatch correction. Proceedings of the National Academy of Sciences of the United States of America96(24), 13914–13919.

Hunter, N., & Borts, R. H. (1997). Mlh1 is unique among mismatch repair proteins in its ability to promote crossing-over during meiosis. Genes & development11(12), 1573–1582.

Chambers, S. R., Hunter, N., Louis, E. J., & Borts, R. H. (1996). The mismatch repair system reduces meiotic homeologous recombination and stimulates recombination-dependent chromosome loss. Molecular and cellular biology16(11), 6110–6120.

Hunter, N., Chambers, S. R., Louis, E. J., & Borts, R. H. (1996). The mismatch repair system contributes to meiotic sterility in an interspecific yeast hybrid. The EMBO journal15(7), 1726–1733.

Németh, A. H., Hunter, N., Coleman, M. P., Borts, R. H., Louis, E. J., & Davies, K. E. (1993). Rescue of a single yeast artificial chromosome from a cotransformation event utilizing segregation at meiosis. Genetic analysis, techniques and applications10(6), 123–127.