Kyle Friend Assistant Professor of Chemistry and Biochemistry

Kyle Friend

Science Addition A427

Professional Positions

Washington and Lee University, Department of Chemistry and Biochemistry

  • Assistant Professor 2014-present
  • Visiting Assistant Professor 2013-2014

Professional Affiliations

  • American Chemical Society 2014-present
  • The RNA Society 2002-present


University of Wisconsin - Madison 2008-13

  • Post-doctorate research
  • Mentor - Judith Kimble
  • Regulation of mRNA translation in C. elegans and mammalian stem cells

Yale University 2007-08

  • Post-doctorate research
  • Mentor - Emre Seli
  • Regulation of the embryonic poly(A)-binding protein

Yale University 2001-07

  • Ph.D. in Molecular Biophysics and Biochemistry
  • Thesis Adviser - Joan Steitz
  • Role of U2 snRNP in intronless mRNA 3' end formation

National Institutes of Health - National Institute of Allergy and Infectious Disease 2000-01

  • NIH Post-baccalaureate IRTA
  • Mentor - Ron Schwartz
  • Molecular mechanism of T cell anergy

Ohio State University 1996-2000

  • B.S. in Biochemistry
  • Mentor - Venkat Gopalan
  • Arabidopsis thaliana RNase P


The Friend lab studies the connections between messenger RNA translation and cell fate decisions. Using mouse embryonic stem cells as a model, we are investigating how mRNA translation is regulated as stem cells differentiate into other cell types, mainly neurons. Our research has shown that key stem cell factors are regulated by mRNA translation and that we can drive the expression of Nanog and c-Myc (two of these factors) by regulating the phosphorylation of a translation initiation factor, eIF2alpha.


  • CHEM 150 - Genetic Engineering and Society
  • CHEM 341 - Biochemistry I
  • CHEM 342 - Biochemistry II
  • CHEM 343 - Biochemistry I Lab
  • CHEM 344 - Biochemistry II Lab

Selected Publications


  • Yoon, D.S., Alfhili, M.A., Friend, K., and Lee, M.H. MPK-1/ERK regulatory network controls the number of sperm by regulating timing of sperm-oocyte switch in C. elegans germline. 2017. Biochem. Biophys. Res. Commun. 491(4): 1077-82. PMID: 28782521.
  • Abebrese, E. L.*, Ali, S.H.*, Andrews, V.M.*, Arnold, Z.R.*, Armstrong, K.A.*, Burns, L.N.*, Crowder, H.R.*, Day Jr., R.T.*, Hsu, D.G.*, Jarrell, K.L.*, Lee, G.M.*, Luo, Y.*, Mugayo, D.A.*, Raza, Z.*, and Friend, K. Identification of human short introns. 2017. PLoS One. 12(5):e0175393. PMID: 28520720.
  • Friend, K., *Brooks, H.A., Propson, N.E., Thomson, J.A. and Kimble, J. 2015. Embryonic stem cell growth factors regulate eIF2α phosphorylation. PLoS One 10(9): e0139076. PMID: 26406898.
  • Friend, K., Brook, M., *Bezirci, F.B., Sheets, M.D., Gray, N.K. and Seli, E. 2012. Embryonic poly(A)-binding protein (ePAB) phosphorylation is required for Xenopus oocyte maturation. Biochem J. 445(1): 93-100. PMID: 22497250.
  • Campbell, Z.T., Menichelli, E., Friend, K., Wu, J., Kimble, J., Williamson, J.R. and Wickens, M. 2012. Identification of a conserved interface between PUF and CPEB proteins. J. Biol. Chem. 287(22): 18854-62. PMID: 22496444.
  • Friend, K., Campbell, Z.T., Cooke, A., Kroll-Conner, P., Wickens, M.P. and Kimble, J. 2012. A conserved PUF—Ago—eEF1A complex attenuates translation elongation. Nat. Struct. Mol. Biol. 19(2): 176-83. PMID: 22231398. Comment in Pasquinelli, A. E. Nat. Struct. Mol. Biol. 19(2): 133-4.
  • Friend, K., Kolev, N.G., Shu, M.D., and Steitz, J.A. 2008. Minor-class splicing occurs in the nucleus of the Xenopus oocyte. RNA 14(8): 1459-1462. PMID: 18567814.
  • Friend, K., *Lovejoy, A.F., and Steitz, J.A. 2007. U2 snRNP binds intronless histone pre-mRNAs to facilitate U7-snRNP-dependent 3' end formation. Mol. Cell 28(2): 240-252. PMID: 17964263. Comment in Marzluff, W.F. Mol. Cell 28(3): 353-354.
  • Fok, V., Friend, K., and Steitz, J.A. 2006. Epstein-Barr virus noncoding RNAs are confined to the nucleus, whereas their partner, the human La protein, undergoes nucleocytoplasmic shuttling. J. Cell. Biol. 173(3): 319-325. PMID: 16682524.
  • Cahill, N., Friend, K., Speckmann, W., Li, Z.H., Terns, R.M., Terns, M.P., Steitz, J.A. 2002. Site-specific cross-linking analyses reveal an asymmetric protein distribution for a box C/D snoRNP. EMBO J. 21(14): 3816-3828. PMID: 12110593.

*denotes undergraduate co-authors.

Book Chapters

  • Kershner, A., Crittenden, S.L., Friend, K., Sorensen, E.B., Porter, D.F., and J. Kimble. 2013. Germline stem cells and their regulation in the nematode Caenorhabditis elegans. In Transcriptional and Translational Regulation of Stem Cells (eds. G. Hime & H. Abud, Springer).
  • Friend, K. and Seli, E. 2009. Molecular biology of the gamete. In Stem Cells in Human Reproduction, 2nd ed. (eds. C. Simon and A. Pellicer, Informa Healthcare Press).


1. An, E. and Friend, K. Investigating ribosome processivity. 2016 Rustbelt RNA Meeting. Cleveland, OH.

2. Walker, A. and Friend, K. Characterization of the RNA-binding motif of Phosphoglycerate Kinase 1 – a Moonlighting Function for a Metabolic Enzyme. 2016. American Society for Biochemistry and Molecular Biology. San Diego, CA.

3. Abebrese, E. L., Ali, S.H., Andrews, V.M., Arnold, Z.R., Armstrong, K.A., Burns, L.N., Crowder, H.R., Day Jr., R.T., Hsu, D.G., Jarrell, K.L., Lee, G.M., Luo, Y., Mugayo, D.A., Raza, Z., and Friend, K. Identification of human short introns. 2015 Gordon Research Conference on Nucleic Acids. Biddeford, ME.

4. Friend, K., Brooks, H.A., Propson, N.E., Thomson, J.A. and Kimble, J. Embryonic stem cells control translation initiation to bypass mRNA upstream open reading frames. 2014 Annual Meeting of the RNA Society. Quebec City, Quebec.

Students Mentored

Fifteen students mentored, nine students at W&L.

Erin An, Gianluca Carrilho-Malta, Bailey Sanders, Billy Davis, Zack Dubit, Austin Walker, Conan Zhao, Emmanuel Abebrese, Daphine Mugayo, Nick Noble, Aalekhya Tenali, Hunter Brooks, Tram Lai (at UW - Madison), Betul Bezirci (at Yale University), Alex Lovejoy (at Yale University)



PI, Jeffress Memorial Trust, 2017-18.
- For investigating how translation is controlled as mouse embryonic stem cells differentiate using a combination of high-throughput sequencing and cell biological assays.


Postdoctoral Fellowship, American Cancer Society, PF-10-127-01-DDC, 2010-13
- Used a combination of in vitro biochemistry and biophysics to characterize a sequence-specific RNA-binding protein. Generated transgenic animals and worked with mammalian cell lines for in vivo functional analysis.