Everett Peter Greenberg
Born (1948-11-07) November 7, 1948 (age 75)[3]
EducationWestern Washington University (BA)
University of Iowa (MSc)
University of Massachusetts Amherst (PhD)
Known forResearch of quorum sensing
SpouseCaroline Harwood
AwardsShaw Prize in Life science and Medicine
Princess of Asturias Award for Technical and Scientific Research
Canada Gairdner International Award
Scientific career
InstitutionsUniversity of Washington
University of Iowa
Cornell University
Harvard University[1]
ThesisThe biology of facultatively anerobic spirochetes (1977)
Doctoral advisorErcole Canale-Parola[2]

Everett Peter Greenberg (born November 7, 1948) is an American microbiologist. He is the inaugural Eugene and Martha Nester Professor of Microbiology at the Department of Microbiology of the University of Washington School of Medicine.[5] He is best known for his research on quorum sensing, and has received multiple awards for his work.

Early life and education

Greenberg was born in 1948 in New York City. When he was an infant, his family moved to San Francisco then to Seattle, where he finished high school.[2][4] He became interested in biology, especially invertebrate biology, after a field trip his biology teacher organized.[4]

Greenberg was determined to study biology after a year at Everett Junior College (now Everett Community College), and, instead of the University of Washington which he thought was too big, opted for Western Washington University in Bellingham, Washington in 1966.[2][4] He graduated with a BA in Biology in 1970.[1]

He discovered his passion for microbiology during his undergraduate years, so despite initially planning to pursue biochemistry for postgraduate study, he eventually went to the University of Iowa to conduct microbiology research, where he obtained his MSc in Microbiology in 1972. Greenberg then started his PhD at the University of Massachusetts Amherst, and received his PhD in 1977.[2]


After obtaining his PhD, Greenberg spent time at Harvard University for postdoctoral work.[2] He joined the Department of Microbiology of Cornell University as an assistant professor in 1979,[2] and was promoted to associate professor in 1984.[1] Greenberg went to the Department of Microbiology of the University of Iowa as a professor in 1988, and returned to Seattle in 2005 when he joined the Department of Microbiology of the University of Washington as head of the department.[4][6]

Outside of academia, Greenberg was an associate editor of the Annual Review of Microbiology from 1991 to 2001.[7]


Before the 1960s and 70s, bacteria were thought to be independent organisms that do not communicate with one another.[8][9] The first indication of inter-bacterial communication was reported by Alexander Tomasz in 1965, who found Streptococcus pneumoniae secretes a substance that allow other S. pneumoniae to take up foreign DNA.[10] Then in 1970, John Woodland Hastings discovered that Aliivibrio fischeri produced light and became bioluminescent under high cell density but not in diluted concentration, a phenomenon known as autoinduction.[11] The molecule that A. fischeri secretes for autoinduction is called an autoinducer, which was not identified until 1981.

In 1985, Greenberg reported that, instead of a complex mechanism, the autoinducer simply passively diffuses from one bacterial cell to another,[12] revealing one of the mechanisms of bacterial communication.

He switched course after moving to the University of Iowa, focusing on the luxR protein in A. fischeri; the protein is a transcription factor activated by the autoinducer.[13] Research by Greenberg's team confirmed the C-terminus of luxR binds DNA[14] while its N-terminus binds the autoinducer.[15]

In a seminal article in 1994, Greenberg, together with Claiborne (Clay) Fuqua and Stephen Winans, at the time both from Cornell University, coined the term quorum sensing to describe the behavior of autoinduced bioluminescence in A. fischeri and other bacterial species.[16]

Greenberg has also branched out from A. fischeri, collaborating with Barbara Iglewski at the University of Rochester to study quorum sensing in Pseudomonas aeruginosa, the infection of which is the primary cause of death in cystic fibrosis patients. The collaboration stemmed from Iglewski's discovery in 1991 of a P. aeruginosa gene coding for a transcription factor protein that controls the expression of genes which, in turn, control its virulence. The closest relative of the transcription factor, in terms of protein sequence, was the luxR protein.[17] This partnership led to the discovery of three quorum sensing signaling pathways in P. aeruginosa.[18]

His research in quorum sensing led Greenberg to study the phenomenon of biofilm, which is a high-density cluster of bacteria that attaches to each other and to surfaces and is embedded in an extracellular matrix. Notably, bacteria in biofilms are more resistant to antibiotics than their free-living counterparts.[13] In 1998, Greenberg, Iglewski, and J. William Costerton reported the link between quorum-sensing genes and biofilm formation, the first publication to show a genetic factor in biofilm structure.[19] Over the next years, he kept studying biofilms and their effects on diseases, discovering that lungs of cystic fibrosis patients are infected with biofilms,[20] and that iron is required for P. aeruginosa biofilm formation.[21]

Personal life

Greenberg met his wife, Caroline Harwood, during his PhD years at the University of Massachusetts Amherst.[4] They started dating when he was a postdoctoral researcher at Harvard University, and were married in 1984.[2] Harwood is currently also a professor at the Department of Microbiology of the University of Washington School of Medicine.[22]

Honors and awards


  1. ^ a b c "Everett Peter Greenberg". Frontiers Media. Archived from the original on August 8, 2023. Retrieved August 8, 2023.
  2. ^ a b c d e f g "Autobiography of E Peter Greenberg". Shaw Prize. Archived from the original on August 9, 2023. Retrieved August 9, 2023.
  3. ^ a b "Jeffrey I. Gordon, Peter Greenberg and Bonnie L. Bassler, Princess of Asturias Award for Technical and Scientific Research". Princess of Asturias Foundation. July 6, 2023. Archived from the original on August 8, 2023. Retrieved August 8, 2023.
  4. ^ a b c d e f Davis, Tinsley H. (2004). "Biography of E. P. Greenberg". Proceedings of the National Academy of Sciences. 101 (45): 15830–15832. doi:10.1073/pnas.0407738101. PMC 528758. PMID 15520366.
  5. ^ "Eavesdropping on microbe chatter earns Gairdner Award". University of Washington School of Medicine. March 29, 2023. Archived from the original on August 9, 2023. Retrieved August 9, 2023.
  6. ^ "Distinguished molecular microbiologist to chair department at UW School of Medicine". University of Washington. October 8, 2004. Archived from the original on August 21, 2023. Retrieved August 21, 2023.
  7. ^ Ornston, Nick (2001). "Preface by Nick Ornston". Annual Review of Microbiology. 55. doi:10.1146/annurev.mi.55.010101.100001.
  8. ^ Greenberg, E. Peter (2003). "Bacterial communication: Tiny teamwork". Nature. 424 (6945): 134. Bibcode:2003Natur.424..134G. doi:10.1038/424134a. PMID 12853935.
  9. ^ Bassler, Bonnie L.; Losick, Richard (2006). "Bacterially Speaking". Cell. 125 (2): 237–246. doi:10.1016/j.cell.2006.04.001. PMID 16630813.
  10. ^ Tomasz, Alexander (1965). "Control of the competent state in Pneumococcus by a hormone-like cell product: an example for a new type of regulatory mechanism in bacteria". Nature. 208 (5006): 155–159. Bibcode:1965Natur.208..155T. doi:10.1038/208155a0. PMID 5884251. S2CID 4202362. Retrieved August 11, 2023.
  11. ^ Nealson, Kenneth H.; Platt, Terry; Hastings, J. Woodland (1970). "Cellular Control of the Synthesis and Activity of the Bacterial Luminescent System". Journal of Bacteriology. 104 (1): 313–322. doi:10.1128/jb.104.1.313-322.1970. PMC 248216. PMID 5473898.
  12. ^ Kaplan, Heidi B.; Greenberg, E. P. (1985). "Diffusion of Autoinducer Is Involved in Regulation of the Vibrio fischeri Luminescence System". Journal of Bacteriology. 163 (3): 1210–1214. doi:10.1128/jb.163.3.1210-1214.1985. PMC 219261. PMID 3897188.
  13. ^ a b Whiteley, Marvin; Diggle, Stephen P.; Greenberg, E. Peter (2017). "Progress in and promise of bacterial quorum sensing research". Nature. 551 (7680): 313–320. Bibcode:2017Natur.551..313W. doi:10.1038/nature24624. PMC 5870893. PMID 29144467.
  14. ^ Choi, S. H.; Greenberg, E. P. (1991). "The C-terminal region of the Vibrio fischeri LuxR protein contains an inducer-independent lux gene activating domain". Proceedings of the National Academy of Sciences. 88 (24): 11115–11119. Bibcode:1991PNAS...8811115C. doi:10.1073/pnas.88.24.11115. PMC 53084. PMID 1763027.
  15. ^ Hanzelka, Brian L.; Greenberg, E. P. (1995). "Evidence that the N-terminal region of the Vibrio fischeri LuxR protein constitutes an autoinducer-binding domain". Journal of Bacteriology. 177 (3): 815–817. doi:10.1128/jb.177.3.815-817.1995. PMC 176662. PMID 7836318.
  16. ^ Fuqua, W. Claiborne; Winans, Stephen C.; Greenberg, E. Peter (1994). "Quorum sensing in bacteria: the LuxR-LuxI family of cell density-responsive transcriptional regulators". Journal of Bacteriology. 176 (2): 269–275. doi:10.1128/jb.176.2.269-275.1994. PMC 205046. PMID 8288518.
  17. ^ Gambello, Michael J.; Iglewski, Barbara H. (1991). "Cloning and characterization of the Pseudomonas aeruginosa lasR gene, a transcriptional activator of elastase expression". Journal of Bacteriology. 173 (9): 3000–3009. doi:10.1128/jb.173.9.3000-3009.1991. PMC 207884. PMID 1902216.
  18. ^ Miranda, Samantha Wellington; Asfahl, Kyle L.; Dandekar, Ajai A.; Greenberg, E. P. (2022). "Pseudomonas aeruginosa Quorum Sensing". In Filloux, Alain; Ramos, Juan-Luis (eds.). Pseudomonas aeruginosa: Biology, Pathogenesis and Control Strategies. Advances in Experimental Medicine and Biology. Vol. 1386. Springer Science+Business Media. pp. 95–115. doi:10.1007/978-3-031-08491-1_4. ISBN 9783031084904. PMC 9942581. PMID 36258070.
  19. ^ Davies, David G.; Parsek, Matthew R.; Pearson, James P.; Iglewski, Barbara H.; Costerton, J. W.; Greenberg, E. P. (1998). "The Involvement of Cell-to-Cell Signals in the Development of a Bacterial Biofilm". Science. 280 (5361): 295–298. Bibcode:1998Sci...280..295D. doi:10.1126/science.280.5361.295. PMID 9535661. Retrieved August 21, 2023.
  20. ^ Singh, Pradeep K.; Schaefer, Amy L.; Parsek, Matthew R.; Moninger, Thomas O.; Welsh, Michael J.; Greenberg, E. P. (2000). "Quorum-sensing signals indicate that cystic fibrosis lungs are infected with bacterial biofilms". Nature. 407 (6805): 762–764. Bibcode:2000Natur.407..762S. doi:10.1038/35037627. PMID 11048725. S2CID 4372096. Retrieved August 21, 2023.
  21. ^ Banin, Ehud; Vasil, Michael L.; Greenberg, E. Peter (2005). "Iron and Pseudomonas aeruginosa biofilm formation". Proceedings of the National Academy of Sciences. 102 (31): 11076–11081. Bibcode:2005PNAS..10211076B. doi:10.1073/pnas.0504266102. PMC 1182440. PMID 16043697.
  22. ^ "Caroline (Carrie) Harwood". Archived from the original on August 10, 2023. Retrieved August 10, 2023.
  23. ^ "Elected Fellows". American Association for the Advancement of Science. Archived from the original on August 18, 2023. Retrieved August 18, 2023.
  24. ^ "Everett Peter Greenberg". American Academy of Arts and Sciences. Archived from the original on August 18, 2023. Retrieved August 18, 2023.
  25. ^ "E. Peter Greenberg". National Academy of Sciences. Archived from the original on August 18, 2023. Retrieved August 18, 2023.
  26. ^ "The 2015 Prize in Life Science & Medicine". Shaw Prize. Archived from the original on August 18, 2023. Retrieved August 18, 2023.
  27. ^ "E Peter Greenberg". Gairdner Foundation. Archived from the original on August 11, 2023. Retrieved August 11, 2023.