Be sure to check out our slidecasts, which are short recorded talks about the work in the papers. It's faster than reading!

If you are interested in getting started with single molecule RNA FISH yourself, check out this website. It has a lot of information and answers to many common questions.

If you want to count your spots, use this software. It is easy to use and gives good results.


Padovan-Merhar O, Nair GP, Biaesch A, Mayer A, Scarfone S, Foley SW, Wu AR, Churchman LS, Singh A, Raj A. Single mammalian cells compensate for differences in cellular volume and DNA copy number through independent global transcriptional mechanisms. Molecular Cell 2015, in press.

Cabili MN*#, Dunagin MC*, McClanahan PD, Biaesch A, Padovan-Merhar O, Regev A*, Rinn JL*#, Raj A*#. Localization and abundance analysis of human lncRNAs at single cell and single molecule resolution. Genome Biology, in press. * equal contributions, # corresponding author. Provisional PDF.


Dimitrova N, Zamudio JR, Jong RM, Soukup D, Resnick R, Sarma K, Ward AJ, Raj A, Lee JT, Sharp PA, Jacks T. LincRNA-p21 Activates p21 In cis to Promote Polycomb Target Gene Expression and to Enforce the G1/S Checkpoint. Molecular Cell 2014, doi:10.1016/j.molcel.2014.04.025 pubmed.

Abranches E, Guedes AM, Moravec M, Maamar H, Svoboda P, Raj A, Henrique D. Stochastic NANOG fluctuations allow mouse embryonic stem cells to explore pluripotency. Development 2014, doi:10.1242/dev.108910 pubmed.

Weis B, Schmidt J, Maamar H, Raj A, Lin H, Tóth C, Riedmann K, Raddatz G, Seitz HK, Ho AD, Lyko F, Linhart HG. Inhibition of intestinal tumor formation by deletion of the DNA methyltransferase 3a. Oncogene 2014, doi:10.1038/onc.2014.114 pubmed.

Ginart P., Raj A. RNA sequencing in situ. Nature Biotechnology 2014 [News and Views], doi:10.1038/nbt.2921 PDF.

Nair, G. An intuitive derivation of the expectation-maximization algorithm. Rajlab, brief communication. PDF.

Facciponte JG, Ugel S, De Sanctis F, Li C, Wang L, Nair G, Sehgal S, Raj A, Matthaiou E, Coukos G, Facciabene A. Tumor endothelial marker 1-specific DNA vaccination targets tumor vasculature. Journal of Clinical Investigation 2014, doi:10.1172/JCI67382. pubmed.

Paralkar VR, Mishra T, Luan J, Yao Y, Kossenkov AV, Anderson SM, Dunagin M, Pimkin M, Gore M, Sun D, Konuthula N, Raj A, An X, Mohandas N, Bodine DM, Hardison RC, Weiss MJ. Lineage and species-specific long noncoding RNAs during erythro-megakaryocytic development. Blood 2014, doi:10.1182/blood-2013-12-544494. pubmed.

Hacisuleyman E, Goff LA, Trapnell C, Williams A, Henao-Mejia J, Sun L, McClanahan P, Hendrickson DG, Sauvageau M, Kelley DR, Morse M, Engreitz J, Lander ES, Guttman M, Lodish HF, Flavell R, Raj A, Rinn JL. Topological organization of multichromosomal regions by the long intergenic noncoding RNA Firre. Nature SMB 2014, doi:10.1038/nsmb.2764. pubmed.


Shaffer, S.M., Wu, M., Levesque, M.J., Raj, A. Turbo FISH: A method for rapid single molecule RNA FISH. PLoS ONE 8(9): e75120, 2013, doi:10.1371/journal.pone.0075120. PDF, protocol, live demo.

Padovan-Merhar, O., Raj, A. Using variability in gene expression as a tool for studying gene regulation. WIREs Syst Biol Med [review article], 2013, doi: 10.1002/wsbm.1243. PDF.

Levesque, M.J., Ginart, P., Wei, Y., Raj, A. Visualizing SNVs to quantify allele-specific expression in single cells. Nature Methods, 2013, doi:10.1038/nmeth.2589. PDF, supplement, slidecast

Nair, G., Walton, T., Murray, J.I.*, Raj, A.* Gene transcription is coordinated with but not dependent on cell divisions during C. elegans embryonic fate specification. Development, 2013, doi:10.1242/dev.098012. (* co-corresponding authors) PDF, supplement, slidecast

Zhang, X., Song, Y., Shah, A.Y., Lekova, V., Raj, A., Huang, L., Behlke, M.A., Tsourkas, A. Quantitative assessment of ratiometric bimolecular beacons as a tool for imaging single engineered RNA transcripts and measuring gene expression in living cells. Nucleic Acids Research, 2013, doi:10.1093/nar/gkt561. link.

Maamar, H., Cabili, N.M., Rinn, J.L., Raj, A. linc-HOXA1 is a non-coding RNA that represses Hoxa1 in cis. Genes and Development, 2013, doi:10.1101/gad.217018.113. PDF, supplement, slidecast

Levesque, MJ, Raj, A Single chromosome transcriptional profiling reveals chromosome-level regulation of gene expression. Nature Methods, 2013, doi:10.1038/nmeth.2372. PDF, raw data+supplement+long version, slidecast

Kretz M, Siprashvili Z, Chu C, Webster DE, Zehnder A, Qu K, Lee CS, Flockhart RJ, Groff AF, Chow J, Johnston D, Kim GE, Spitale RC, Flynn RA, Zheng GX, Aiyer S, Raj A, Rinn JL, Chang HY, Khavari PA. Control of somatic tissue differentiation by the long non-coding RNA TINCR. Nature, 2013, doi:10.1038/nature11661. Link


Mirouze, N., Desai, Y., Raj, A., Dubnau, D. Spo0A-P imposes a temporal gate for the bimodal expression of competence in B. subtilis. PLoS Genetics, 2012, doi:10.1371/journal.pgen.1002586. PDF

Li, Fan; Zheng, Qi; Ryvkin, Paul; Dragomir, Isabelle; Desai, Yaanik; Aiyer, Subhadra; Valladares, Otto; Yang, Jamie; Bambina, Shelly; Sabin, Leah R.; Murray, John I.; Lamitina, Todd; Raj, Arjun; Cherry, Sara; Wang, Li-San; Gregory, Brian D. Global Analysis of RNA Secondary Structure in Two Metazoans. Cell Reports 2012; 1:69-82. PDF with Supplementary Information.


Nair G, Raj A. Time-lapse transcription. Science [perspective]. 2011 Apr 22; 332(6028):431-2. PDF.

Steine EJ, Ehrich M, Bell GW, Raj A, Reddy S, van Oudenaarden A, Jaenisch R, Linhart HG. Genes methylated by DNA methyltransferase 3b are similar in mouse intestine and human colon cancer. J Clin Invest. 2011 May 2;121(5):1748-52. doi: 10.1172/JCI43169. PDF, Supplementary Information.

Batish M, Raj A, Tyagi S. Single molecule imaging of RNA in situ. Methods Mol Biol. 2011;714:3-13. PDF.


Raj A, Tyagi S. Detection of individual endogenous RNA transcripts in situ using multiple singly labeled probes. Method in Enzymology 2010; 472: 365-86. PDF.

Raj A*, Rifkin SA*, Andersen E, van Oudenaarden A. Variability in gene expression underlies incomplete penetrance. Nature, Feb 18 2010, doi:10.1038/nature08781 (* equal contributions). PDF, Supplementary Information, Data, Slidecast, News and Views, NPR, Cell Preview, NRC Handelsblad, Nat Rev Gen, Word cloud.


Khalil AM, Guttman M, Huarte M, Garber M, Raj A, Rivea Morales D, Thomas K, Presser A, Bernstein BE, van Oudenaarden A, Regev A, Lander ES, Rinn JL. Many human large intergenic noncoding RNAs associate with chromatin-modifying complexes and affect gene expression. PNAS 2009; 106: 11667-72. PDF.

Raj A, van Oudenaarden A. Single-Molecule Approaches to Stochastic Gene Expression [Review article]. Ann. Rev. Biophysics 2009; 38: 255-70. PDF.


Raj A, van Oudenaarden A. Nature, Nurture or Chance: Stochastic Gene Expression and its Consequences [Review article]. Cell 2008; 135(2): 216-226. PDF.

Raj A, van den Boogard P, Rifkin SA, van Oudenaarden A, Tyagi S. Imaging individual mRNA molecules using multiple singly labeled probes. Nature Methods 2008; 5(10): 877-879. PDF, Supplementary material, Software.

To design your own probes, visit


Maamar H*, Raj A*, Dubnau D. Noise in Gene Expression Determines Cell Fate in Bacillus Subtilis. Science, 2007 Jul 23; 317(5837): 526-9 (* equal contributions). PDF, Supplementary material, Movies

Raj A, Peskin CS, Tranchina D, Vargas DY, Tyagi S. Stochastic mRNA Synthesis in Mammalian Cells. PLoS Biol 2006; 4(10): e309. PDF, Supplementary material, Data

Raj A, Peskin CS. The influence of chromosome flexibility on chromosome transport during anaphase A. Proc Natl Acad Sci USA, 2006 Apr 4;103(14): 5349-54. PDF, Supplementary material

Vargas DY, Raj A, Marras SA, Kramer FR, Tyagi S. Mechanism of mRNA transport in the nucleus. Proc Natl Acad Sci USA, 2005 Nov 22;102(47): 17008-13. PDF, Supplementary material

Man, I'm getting old...

Raj, A., T. Phan, R. P. Lin, and V. Angelopoulos, Wind survey of high-speed bulk flows and field-aligned beams in the near-Earth plasma sheet. J. Geophys. Res. 2002, 107(A12), 1419, PDF.

Gibbons, M. Raj, A. Strichartz, R. S. The Finite Element Method on the Sierpinski Gasket. Constructive Approximation, 2001, 17(4): 561-588, PDF, website.