Dr. Irvin Pan earned his B.A. in Biology from Amherst College, where he did undergraduate research in ecology. He then attended the Writing Seminars at Johns Hopkins University, where he earned an M.A. in Science Writing. Dr. Pan earned his Ph.D. in Molecular, Cellular, and Developmental Biology from Yale University. At Yale, his work in the lab of Dr. Vivian Irish focused on examining the function and evolution of a transcription factor involved in tomato fruit development and ripening. Prior to coming to Stonehill College, Dr. Pan was a postdoctoral associate and a TEACRS (Training in Education and Critical Research Skills) Fellow at Tufts University in the lab of Dr. John Coffin, where he studied the evolution of a family of mouse endogenous retroviruses.

Dr. Pan’s research focuses on a group of MADS box transcription factors that have acquired distinct and novel functions in controlling fruit development in Arabidopsis and tomato. He plans to further examine the developmental pathways that these genes control to unravel the genetic changes that have lead to the evolution and domestication of the modern day tomato.

Recent Student Projects

  • The role of Tomato Agamous-Like 1 (TAGL1) in fruit development in tomato and other related species
  • Identification and characterization of cold-tolerant crop plants for four-season harvesting


  • B.A., 1999, Biology, Amherst College
  • M.A., 2000, Science Writing, Johns Hopkins University
  • Ph.D., 2008, Molecular, Cellular, and Developmental Biology, Yale University

Research Interests

  • Plant evolutionary developmental biology
  • The genetics of domesticated crops
  • Viral evolution

Courses Taught

  • Developmental Biology
  • Biological Principles I
  • Biological Principles II
  • Virology
  • Genetics Laboratory

Areas of Expertise


Associate Professor of Biology



  • Pan, I.L., McQuinn, R., Giovannoni, J.J., and Irish, V.F. 2010. Functional diversification of AGAMOUS lineage genes in regulating tomato flower and fruit development. Journal of Experimental Botany. 61, 1795-1806.
  • Vrebalov, J.*, Pan, I.L.*, Arroyo, A.J., McQuinn, R., Chung, M., Poole, M., Rose, J., Seymour, G.,Grandillo, S., Giovannoni, J., and Irish, V.F. 2009. Fleshy fruit expansion and ripening are regulated by the tomato SHATTERPROOF gene TAGL1. Plant Cell 21: 3041-3062. *Co-first authors
  • De Martino, G., Pan, I., Emmanuel, E., Levy, A.., Irish, V.F. 2006. Functional analyses of two tomato APETALA3 genes demonstrate diversification in their roles in regulating floral development. Plant Cell 18: 1833-1845.
  • Van Dover et al. 2001. Biogeography and ecological setting of Indian Ocean hydrothermal vents. Science 294: 818-823.
  • Temeles, E.J., Pan, I.L., Brennan, J.L, Horwitt, J.N. 2000. Evidence for ecological causation of sexual dimorphism in a hummingbird. Science 289: 441-443.