2021 – present

38. Bree Heestand, Ben McCarthy, Matt Simon, Evan-Lister Shimauchi, Stephen Frenk, Shawn Ahmed (2024). Deficiency for Piwi results in transmission of a heritable stress that promotes longevity via DAF-16/Foxo. Manuscript resubmitted to Aging Cell. bioRxiv. 2024 June 24 [Preprint].

37. Evan H. Lister-Shimauchi, Morgan Brady , Stephen Frenk, Braxton Harris, Anna-Maria Leon Ortiz, Aylin Memili, Minh Nguyen, Simon Boulton, Shawn Ahmed (2024). RTEL-1 and DNA polymerase theta promote subtelomeric DNA synthesis and telomere fusion in C. elegans. Manuscript in revision at eLife. bioRxiv. 2024 Feb 4 [Preprint].

36. Peter Mark Carlton, Richard E. Davis, Shawn Ahmed (2022). Nematode chromosomes. Genetics 2022 Mar 22:iyac014. doi: 10.1093/genetics/iyac014. PMID: 35323874

35. Evan H. Lister-Shimauchi, Benjamin McCarthy, Michael Lippincott, Shawn Ahmed. Genetic and Epigenetic Inheritance at Telomeres. Epigenomes 2022 Mar; 6(1): 9. Published online 2022 Mar 16. doi: 10.3390/epigenomes6010009

34. Evan H. Lister-Shimauchi, Michael Dinh, Paul Maddox, Shawn Ahmed (2021). Gametes deficient for Pot1 telomere binding proteins alter levels of telomeric foci for multiple generations. Communications Biology Feb 4;4(1):158. doi: 10.1038/s42003-020-01624-7. PubMed PMID: 33542458; PubMed Central PMCID: PMC7862594

33. Maya Spichal, Bree Heestand, Katherine Kretovich Billmyre, Stephen Frenk, Craig C. Mello, Shawn Ahmed (2021). Germ granule dysfunction is a hallmark and mirror of Piwi mutant sterility. Nature Communcations Mar 3;12(1):1420. doi: 10.1038/s41467-021-21635-0. PubMed PMID: 33658512; PubMed Central PMCID: PMC7930041

2011 – 2020

32. Stephen Frenk, Evan Lister-Shimauchi and Shawn Ahmed (2019). Telomeric small RNAs in the genus Caenorhabditis. RNA  25: 1061-1077. PMID: 31239299

31. Katherine Kretovich Billmyre, Anna-Lisa Doebley, Bree Heestand, Tony Belicard, Aya Sato-Carlton, Stephane Flibotte, Matt Simon, Megan Gnazzo, Ahna Skop, Donald Moerman, Peter Mark Carlton, Peter Sarkies and Shawn Ahmed (2019). The conserved phosphatase GSP-2/PP1 promotes germline immortality via small RNA-mediated genome silencing during meiosis. PLOS Genetics

30. Bree Heestand, Matt Simon, Stephen Frenk, Denis Titov and Shawn Ahmed (2018). Transgenerational sterility of Piwi mutants represents a dynamic form of Adult Reproductive Diapause. Cell Reports 23:156-171. Available from:

29. Luciana Leopold, Bree Heestand, Soobin Seong, Ludmilla Shtessel and Shawn Ahmed (2015). Lack of pairing during meiosis triggers multigenerational transgene silencing in Caenorhabditis elegans. PNAS  112:E2667-2676. PMCID: PMC4443339

28. Aisa Sakaguchi, Peter Sarkies, Matt Simon, Anna-Lisa Doebley, Leonard D. Goldstein, Ashley Hedges, Kohta Ikegami, Stacy M. Alvares, Liwei Yang, Jeannine R. LaRocque, Julie Hall, Eric A. Miska*, Shawn Ahmed* (2014). C. elegans RSD-2 and RSD-6 promote germ cell immortality by maintaining small interfering RNA populations. PNAS 111:E4323-4331.

*equal contribution

27. Matt Simon, Peter Sarkies, Kohta Ikegami, Anna-Lisa Doebley, Leonard D. Goldstein, Jacinth Mitchell, Aisa Sakaguchi, Eric A. Miska* and Shawn Ahmed* (2014). Reduced insulin/IGF-1 signaling restores germ cell immortality to Caenorhabditis elegans Piwi mutants. Cell Reports 7: 1-12.

*equal contribution

26. Stacy M. Alvares, Gaea A. McCaig, Ebony Y. Joyner, Bernard Lakowski, Shawn Ahmed(2014). H3K4 demethylase activities repress proliferative and post-mitotic aging. Aging Cell, e-pub ahead of print, PMID:24134677

25. Ludmila Shtessel, Mia Lowden, Chen Cheng, Matt Simon, Kyle Wang and Shawn Ahmed (2013). Caenorhabditis elegans POT-1 and POT-2 repress telomere maintenance pathways. G3 3: 305-313. PMID: 23390606

24. Alyson Ashe, Alexandra Sapetschnig, Eva-Maria Weick, Jacinth Mitchell, Marloes P. Bagijn, Amy C. Cording, Anna-Lisa Doebley, Leonard D. Goldstein, Nicolas J. Lehrbach, Jérémie Le Pen, Greta Pintacuda, Aisa Sakaguchi, Peter Sarkies, Shawn Ahmed* and Eric A. Miska* (2012). piRNAs can trigger a multigenerational epigenetic memory in the germline of C. elegans. Cell 150: 88-99. PMID: 22738725

*equal contribution

23. Chen Cheng, Lumilla Shtessel, Megan Brady and Shawn Ahmed (2012). Caenorhabditis elegans POT-2 represses a mode of ALT with normal telomere lengths. PNAS 109: 7805-7810. PMID: 22547822

22. Mia Rochelle Lowden, Stephane Flibotte, Donald Moerman, and Shawn Ahmed (2011). DNA synthesis generates terminal duplications that seal end-to-end chromosome fusions. Science 332: 468-471.

21. Robert H. Dowen and Shawn Ahmed (2019). Maternal Inheritance: Longevity programs nourish progeny via yolk. Current Biology 29(15):R748-R751. doi: 10.1016/j.cub.2019.06.050

20. Ludmila Shtessel and Shawn Ahmed (2011). Telomere dysfunction in human bone marrow failure syndromes. Nucleus 2: 24-29.

2001 – 2010

19. Aymeric P. Bailly, Alasdair Freeman, Julie Hall, Anne-Cécile Déclais, Arno Alpi, David M.J. Lilley, Shawn Ahmed and Anton Gartner (2010). The Caenorhabditis elegans homolog of Gen1/Yen1 resolvases links DNA damage signaling to DNA double-strand break repair. PLoS Genetics 6: e1001025.

18. Bettina Meier, Louise J. Barber, Yan Liu, Ludmila Shtessel, Simon J. Boulton, Anton Gartner* and Shawn Ahmed* (2009). The MRT-1 nuclease is required for DNA crosslink repair and telomerase activity in vivo in C. elegans. EMBO Journal 28: 3549-3563.

*equal contribution

17. Sebastien Greiss, Julie Hall, Shawn Ahmed and Anton Gartner (2008). C. elegans SIR-2.1 translocation is linked to a pro-apoptotic pathway parallel to cep-1/p53 during DNA damage induced apoptosis. Genes and Development 22: 2831-2842.

16. Mia Lowden, Bettina Meier, Teresa Lee, Julie Hall and Shawn Ahmed (2008). End-joining at Caenorhabditis elegans telomeres. Genetics 180: 741-754.

15. Julie Boerckel, Dana Walker and Shawn Ahmed (2007). The RFC-like subunit HPR-17 is required for telomere replication in C. elegans. Genetics 176: 703-709.

14. Shawn Ahmed (2006). Uncoupling of pathways that promote post-mitotic lifespan and apoptosis from replicative immortality of C. elegans germ cells. Aging Cell 5: 559-563.

13. Jasper Harris, Mia Lowden, Iuval Clejan, Monika Tzoneva, James Thomas, Jonathan Hodgkin and Shawn Ahmed (2006). Mutator phenotype of C. elegans DNA damage checkpoint mutants. Genetics 174: 1-16. (Article Highlighted in October issue of Genetics)

12. Iuval Clejan, Julie Boerckel and Shawn Ahmed (2006). Developmental modulation of non-homologous end-joining in C. elegans. Genetics 173: 1301-1317.

11. Bettina Meier, Iuval Clejan, Yan Liu, Mia Lowden, Anton Gartner, Jonathan Hodgkin, and Shawn Ahmed (2006). trt-1 is the C. elegans catalytic subunit of telomerase. PLoS Genetics 2: e18.

10. Jacapo Novelli, Shawn Ahmed and Jonathan Hodgkin (2004). Gene interactions in Caenorhabditis elegans define DPY-31 as a candidate procollagen C-proteinase and SQT-3/ROL-4 as its predicted major target. Genetics 168: 1259-1273.

9. Bettina Meier and Shawn Ahmed (2001). Checkpoints: Meiotic chromosome pairing takes an unexpected twist. Current Biology 11: R865-R868.

8. Shawn Ahmed, Arno Alpi, Michael O. Hengartner and Anton Gartner (2001). C. elegans RAD-5/CLK-2 defines a new DNA damage checkpoint protein. Current Biology 11: 1934-1944.

Review Articles:

7. Theresa Zucchero and Shawn Ahmed (2006). Genetics of proliferative aging. Experimental Gerontology 41: 992-1000. 

6. Chris Smelick and Shawn Ahmed (2005). Achieving immortality in the C. elegans germline. Aging Research Reviews 4: 67-82.

Prior to 2000

5. Anton Gartner, Stuart Milstein, Shawn Ahmed, Jonathan Hodgkin and Michael O. Hengartner (2000). A conserved checkpoint pathway mediates DNA damage-induced apoptosis and cell cycle arrest in C. elegans. Molecular Cell 5: 435-443.

4. Shawn Ahmed and Jonathan Hodgkin (2000). MRT-2 checkpoint protein is required for germline immortality and telomere replication in C. elegans. Nature 403: 159-164.

3. Shawn Ahmed, Hong Sheng, Luming Niu and Eric Henderson (1998). Tetrahymena mutants with short telomeres. Genetics 150: 643-650.

2. Shawn Ahmed, Augustin Kintanar and Eric Henderson (1994). Human telomeric C-strand tetraplexes. Nature Structural Biology 1: 83-88.

1. Shawn Ahmed and Eric Henderson (1992). Formation of novel hairpin structures by telomeric C-strand oligonucleotides. Nucleic Acids Research 20: 507-511.