Steven Irvine


The Irvine Lab is focused on how the regulation of genes affects development and physiology. Recently we have begun to investigate how increases in ocean water temperature may be affecting reproduction through its effect on gene expression. We are also interested in how developmental genes regulate embryogenesis, and how gene regulation has evolved to generate new developmental patterns. Our main study organism is the sea squirt Ciona intestinalis, for which there is an abundance of resources for analyzing gene expression, development and physiology. For more information, visit the Irvine Lab website.


Ph. D. Evolutionary Biology, 1998, University of Chicago
M. Arch. (Architecture), 1983, Harvard University
B. S. Architectural Studies, 1979, University of Illinois

Selected Publications

  1. Irvine SQ (2019) Embryonic canalization and its limits – a view from temperature. In revision.
  2. Irvine SQ, McNulty KB, Siler EM, Jacobson RE (2019) High temperature limits on developmental canalization in the ascidian Ciona intestinalis. Mech. Dev., 157:10-21, 
  3. Irvine SQ, Ristoratore F, Di Gregorio A (2019) Tunicates: From Humble Sea Squirt to Proud Model Organism. Dev Biol, 448: 69-70.
  4. Lopez CE, Sheehan HE, Vierra DA, Azzinaro PA, Meedel TH, Howlett NG, Irvine SQ (2017) Proteomic responses to elevated ocean temperature in ovaries of the ascidian Ciona intestinalis. Biology Open, in press.
  5. Stanley EC, Azzinaro PA, Vierra DA, Howlett NG, Irvine SQ (2016) The simple chordate Ciona intestinalis has a reduced complement of genes associated with Fanconi Anemia. Evolutionary Bioinformatics, 2016:12, 133–148, doi: 10.4137/EBO.S37920.
  6. Irvine SQ (2013) Study of cis-regulatory elements in the ascidian Ciona intestinalis. Curr Genomics 14:56-67.
  7. Vierra DA, Irvine SQ. (2012) Optimized conditions for transgenesis of the ascidian Ciona using square wave electroporation. Dev. Genes Evol. 222:55-61.
  8. Irvine SQ, Vierra DA, Millette BJ, Blanchette MD, Holbert RE (2011) Expression of the Distalless-B gene in Ciona is regulated by a pan-ectodermal enhancer module. Dev. Biol. 344: 457-458.
  9. Irvine SQ, Fonseca VC, Antony R, Zompa MA (2008) Cis-regulation of the Pax6 gene in the ascidian Ciona intestinalis. Developmental Biology 317:649-659.
  10. Irvine SQ, Cangiano MC, Millette BJ, Gutter ES (2007) Non-overlapping expression patterns of the clustered Dll-A/B genes in the ascidian Ciona intestinalis. J. Exp. Zool. (Mol. Dev. Evol.) 308B: 428-441.
  11. Irvine SQ (2007) Whole-mount in-situ hybridization of small invertebrate embryos using laboratory mini-columns. Biotechniques 43:764-768.
  12. Irvine SQ, Seaver EC (2006) Early Annelid Development, A Molecular Perspective. Pp 91-138 in Rouse G, Pleijel F, eds. Reproductive Biology and Phylogeny of Annelida. Jamieson BJM, series ed. Science Publishers: New Hampshire, USA.
  13. Sumiyama K, Irvine SQ, Ruddle FH (2003) The role of gene duplication in the evolution and function of the vertebrate Dlx/distal-less bigene clusters. J. Struc. Func. Genomics 3: 151-159.
  14. Sumiyama K, Irvine SQ, Stock DW, Weiss KM, Kawasaki K, Shimizu N, Shashikant CS, Miller W, Ruddle FH (2002) Genomic structure and functional control of the Dlx3-7 bigene cluster. Proc. Natl. Acad. Sci. USA 99:780-785.
  15. Irvine SQ, Carr JL, Bailey W, Amemiya C, Ruddle FH (2002) Genomic analysis of Hox clusters in the sea lamprey Petromyzon marinus. J. Exp. Zool. (Mol. Dev. Evol.) 294:47-62.
  16. Irvine SQ, Martindale MQ (2001) Comparative analysis of Hox gene expression patterns in the polychaete Chaetopterus: implications for the evolution of body plan regionalization. Am. Zool. 41:640-651.
  17. Irvine SQ, Martindale MQ (2000) Expression patterns of anterior Hox genes in the polychaete Chaetopterus: Correlation with morphological boundaries. Dev. Biol. 217:333-351.
  18. Irvine SQ, Warriner S, Hunter J, Martindale MQ (1997) A survey of homeobox genes in Chaetopterus variopedatus and analysis of polychaete homeodomains. Mol. Phylogen. Evol. 7:331-345.
  19. Panganiban G, Irvine SM, Lowe C, Roehl H, Corley LS, Sherbon B, Grenier JK, Fallon JF, Kimble J, Walker M, Wray GA, Swalla BJ, Martindale MQ, Carroll SB (1997) The origin and evolution of animal appendages. Proc. Natl. Acad. Sci. USA 94:5162-5166. (SMI is co-first author)


BIO 101, Principles of Biology I
BIO/GEO 272, Principles of Evolution
BIO 302, Animal Development
BIO/GEO 572, Advanced Evolutionary Biology