Cohen AA, Ferrucci L, Fülöp T, Gravel D, Hao N, Kriete A, Levine ME, Lipsitz LA, Olde Rikkert MGM, Rutenberg A, Stroustrup N, Varadhan R. A complex systems approach to aging biology. Nat Aging 2022 July; 2:580-591. (pdf)


Paxman J, Zhou Z, O'Laughlin R, Li Y, Tian W, Su H, Jiang Y, Holness SE, Stasiowski E, Tsimring LS, Pillus L, Hasty J, Hao N. Age-dependent aggregation of ribosomal RNA-binding proteins links deterioration in chromatin stability with loss of proteostasis. bioRxiv 2021. (full text)

Choi J, Wang S, Li Y, Hao N, Zid BM. Age-induced P-bodies become detrimental and shorten the lifespan of yeast. bioRxiv 2021. (full text)

Jiang Y, Hao N. Memorizing environmental signals through feedback and feedforward loops. Curr Opin Cell Biol. 2021 April; 69:96-102. (pdf)


Mudla A, Jiang Y, Arimoto K, Xu B, Rajesh A, Ryan AP, Wang W, Daugherty MD, Zhang DE, Hao N. Cell-cycle-gated feedback control mediates desensitization to interferon stimulation. eLife 2020 Sep 18;9:e58825. (pdf) (UCSD News, ScienceDaily, Agile Capital Markets, The Diet World)

Li Y, Jiang Y, Paxman J, O'Laughlin R, Klepin S, Zhu Y, Pillus L, Tsimring LS, Hasty J, Hao N. A programmable fate decision landscape underlies single-cell aging in yeast. Science 2020 July 17; 369(6501):325-329. (pdf) (CNN, Daily Mail UK, Fox 5, UCSD News, KPBS News, Metro World News, GEN, Inverse, PHYS.ORG, Medical News, ScienceDaily, Patch,Technology Networks, Sky tg24 Italy, Sputnik News France, Express Informer)

Jiang Y, AkhavanAghdam Z, Li Y, Zid BM, Hao N. A protein kinase A-regulated network encodes short- and long-lived cellular memories. Sci Signal. 2020 May 19;13(632):eaay3585. (pdf)

O'Laughlin R, Jin M, Li Y, Pillus L, Tsimring LS, Hasty J, Hao N. Advances in quantitative biology methods for studying replicative aging in Saccharomyces cerevisiae. Transl Med Aging. 2020; 4:151-160. (pdf)


Shellhammer JP, Pomeroy AE, Li Y, Dujmusic L, Elston TC, Hao N, Dohlman HG. Quantitative analysis of the yeast pheromone pathway. Yeast. 2019 Apr 25. (pdf)

Jin M, Li Y, O'Laughlin R, Bittihn P, Pillus L, Tsimring LS*, Hasty J*, Hao N*. Divergent aging of isogenic yeast cells revealed through single-cell phenotypic dynamics. Cell Syst. 2019 Mar;8,1-12. (pdf) (* co-correspondence)


Baumgartner B, O'Laughlin R, Jin M, Tsimring LS, Hao N, Hasty J. Flavin-based metabolic cycles are integral features of growth and division in single yeast cells. Sci Rep. 2018 Dec;8:18045. (pdf)


Li Y, Roberts J, AkhavanAghdam Z, Hao N. Mitogen-activated protein kinase (MAPK) dynamics determine cell fate in the yeast mating response. J Biol Chem. 2017 Dec; 292(50):20354-20361. (pdf)

Li Y, Jin M, O'Laughlin R, Bittihn P, Tsimring LS, Pillus L, Hasty J, Hao N. Multigenerational silencing dynamics control cell aging. Proc Natl Acad Sci U S A. 2017 Oct; 114 (42): 11253-11258. (pdf) (UCSD News, KUSI News, abc 10 News, PHYS.ORG, ScienceDaily, Medindia, Kurzweilai, UCSD Guardian)

Shao B, Yuan H, Zhang R, Wang X, Zhang S, Ouyang Q, Hao N*, Luo C*. Reconstructing the regulatory circuit of cell fate determination in yeast mating response. PLoS Comput Biol. 2017 Jul; 13(7):e1005671. (pdf) (* co-correspondence)

Jiang Y, AkhavanAghdam Z,Tsimring LS, Hao N. Coupled feedback loops control the stimulus-dependent dynamics of the yeast transcription factor Msn2. J Biol Chem. 2017 Jun; 292(30):12366-12372. (pdf)

Zhang R, Yuan H, Wang S, Ouyang Q, Chen Y, Hao N, Luo C. High-throughput single-cell analysis for the proteomic dynamics study of the yeast osmotic stress response. Sci Rep. 2017 Feb;7:42200. (pdf)


AkhavanAghdam Z, Sinha J, Tabbaa OP, Hao N. Dynamic control of gene regulatory logic by seemingly redundant transcription factors. eLife 2016 Sep;5. (pdf)

Yuan H, Zhang R, Shao B, Wang X, Ouyang Q, Hao N, Luo C. Protein expression patterns of the yeast mating response. Integr Biol. 2016 Jun;8(6):712-9. (pdf)


Hansen AS, Hao N, O’Shea EK. High-throughput microfluidics to control and measure signaling dynamics in single yeast cells. Nat Protoc. 2015 Aug;10(8):1181-97. (pdf)

2013 and before

Hao N, Budnik BA, Gunawardena J, O’Shea EK. Tunable signal processing through modular control of transcription factor translocation. Science 2013 Jan 25; 339(6118):460-4.

Hao N, Yildirim N, Nagiec MJ, Parnell SC, Errede B, Dohlman HG, Elston TC. Combined computational and experimental analysis reveals MAP kinase-mediated feedback phosphorylation as a mechanism for signaling specificity. Mol Biol Cell. 2012 Oct;23(19):3899-910.

Hao N, O'Shea EK. Signal-dependent dynamics of transcription factor translocation controls gene expression. Nat Struct Mol Biol. 2011 Dec 18;19(1):31-9.

Hao N, Zeng Y, Elston TC, Dohlman HG. Control of MAPK specificity by feedback phosphorylation of shared adaptor protein Ste50. J Biol Chem. 2008 Dec 5;283(49):33798-802.

Behar M, Hao N, Dohlman HG, Elston TC. Dose-to-duration encoding and signaling beyond saturation in intracellular signaling networks. PLoS Comput Biol. 2008 Oct;4(10):e1000197.

Hao N, Nayak S, Behar M, Shanks RH, Nagiec MJ, Errede B, Hasty J, Elston TC, Dohlman HG. Regulation of cell signaling dynamics by the protein kinase-scaffold Ste5. Mol Cell. 2008 Jun 6;30(5):649-56.

Behar M, Hao N, Dohlman HG, Elston TC. Mathematical and computational analysis of adaptation via feedback inhibition in signal transduction pathways. Biophys J. 2007 Aug 1;93(3):806-21.

Hao N, Behar M, Elston TC, Dohlman HG. Systems biology analysis of G protein and MAP kinase signaling in yeast. Oncogene 2007 May 14;26(22):3254-66. Review.

Hao N, Behar M, Parnell SC, Torres MP, Borchers CH, Elston TC, Dohlman HG. A systems-biology analysis of feedback inhibition in the Sho1 osmotic-stress-response pathway. Curr Biol. 2007 Apr 17;17(8):659-67.

Chasse SA, Flanary P, Parnell SC, Hao N, Cha JY, Siderovski DP, Dohlman HG. Genome-scale analysis reveals Sst2 as the principal regulator of mating pheromone signaling in the yeast Saccharomyces cerevisiae. Eukaryot Cell 2006 Feb;5(2):330-46.

Wang X, Hao N, Dohlman HG, Elston TC. Bistability, stochasticity, and oscillations in the mitogen-activated protein kinase cascade. Biophys J. 2006 Mar 15;90(6):1961-78.

Yildirim N, Hao N, Dohlman HG, Elston TC. Mathematical modeling of RGS and G-protein regulation in yeast. Methods Enzymol. 2004;389:383-98. Review.

Hao N, Yildirim N, Wang Y, Elston TC, Dohlman HG. Regulators of G protein signaling and transient activation of signaling: experimental and computational analysis reveals negative and positive feedback controls on G protein activity. J Biol Chem. 2003 Nov 21;278(47):46506-15.