Sean B. Carroll

Sean B. Carroll (born September 17, 1960) is an American evolutionary developmental biologist, author, educator and executive producer. He is the Allan Wilson Professor of Molecular Biology and Genetics at the University of Wisconsin–Madison. His studies focus on the evolution of cis-regulatory elements in the regulation of gene expression in the context of biological development, using Drosophila as a model system. He is a member of the National Academy of Sciences, of the American Philosophical Society (2007),[1] of the American Academy of Arts and Sciences and the American Association for Advancement of Science, as well as a Howard Hughes Medical Institute investigator.

Sean B. Carroll
Carroll in 2008
Born (1960-09-17) September 17, 1960
ResidenceMadison, Wisconsin and Bethesda, Maryland
CitizenshipUnited States
Alma materWashington University in St. Louis (B.S.), Tufts University (Ph.D.)
AwardsPresidential Young Investigator Award
Benjamin Franklin Medal in Life Science
Stephen Jay Gould Prize from the Society for the Study of Evolution
Distinguished Service Award from the National Association of Biology Teachers
Shaw Scientist Award from the Greater Milwaukee Foundation
Scientific career
FieldsEvolutionary developmental biology, Molecular Biology, Genetics
InstitutionsUniversity of Wisconsin–Madison, University of Colorado at Boulder
Doctoral advisorB. David Stollar
Other academic advisorsMatthew P. Scott


Carroll was born in Toledo, Ohio. He has stated that, as a kid, he would flip over rocks looking for snakes, and at age 11 or 12, he started keeping snakes. This activity led him to notice the patterns on the snakes and wonder how those form. Carroll got his B.A. in Biology at Washington University in St. Louis, his Ph.D. in immunology from Tufts University and did post-doctoral work at the University of Colorado Boulder.[2]


Carroll is at the forefront of a field known as evolutionary developmental biology (also known as "evo-devo"), studying how gene changes control the evolution of body parts and patterns. He is the Allan Wilson Professor of Molecular Biology and Genetics at the University of Wisconsin–Madison, and an investigator for the Howard Hughes Medical Institute.[3]

In 1987, Carroll set up a laboratory at the University of Wisconsin-Madison "focused on understanding how genes get used in different ways to generate the diversity of form that we see".[2] The Laboratory of Cell and Molecular Biology lists Carroll's interests as "Genetic control of body pattern in fruit flies, butterflies, and other animals".[4]

Carroll's team has shown, in a series of papers, how the activation of genes during the embryonic stages of the Drosophila fruit fly control the development of its wings, and has been searching for the butterfly's counterparts of these genes.[5]

In 1989, he received the Shaw Scientist Award from the Greater Milwaukee Foundation.[6]

In 2006, Carroll was interviewed by PBS as part of the NOVA documentary "The Family That Walks on All Fours",[7] about a family in Turkey that has members who walk on their hands and feet. In this interview, he discusses the possible genetic underpinnings of this family's condition.

From September 2009[8] to March 2013,[9] he wrote a column for The New York Times called "Remarkable Creatures", where he would discuss findings in animal evolution.

In 2010, he was named vice-president for science education of the Howard Hughes Medical Institute.[10] In 2011, the HHMI launched a documentary film initiative to produce science features for television, to which Carroll was appointed as one of the executive producers.[11] In 2012, one such film, called The Day the Mesozoic Died, retracing the investigation that led to the discovery of the asteroid collision that triggered the mass extinction at the end of that Era, was introduced by Carroll at a National Teacher's Conference.[12]

In 2010, Carroll received the Stephen Jay Gould Prize from the Society for the Study of Evolution.[13][14] In 2012, he was awarded the Benjamin Franklin Medal in Life Science from the Franklin Institute "for proposing and demonstrating that the diversity and multiplicity of animal life is largely due to the different ways that the same genes are regulated rather than to mutation of the genes themselves."[2] In 2016, he was awarded the Lewis Thomas Prize at the Rockefeller University.[15]

Carroll is a proponent of the extended evolutionary synthesis.[16]

Since 2013, Carroll has been listed on the Advisory Council of the National Center for Science Education.[17]

Selected works


  • From DNA to Diversity: Molecular Genetics and the Evolution of Animal Design, with Jennifer Grenier and Scott Weatherbee (2004, Wiley-Blackwell; ISBN 1-4051-1950-0)
  • Endless Forms Most Beautiful: The New Science of Evo Devo and the Making of the Animal Kingdom (2005, W. W. Norton & Company; ISBN 0-393-06016-0)
  • The Making of the Fittest: DNA and the Ultimate Forensic Record of Evolution (2006, W. W. Norton & Company; ISBN 0-393-06163-9)
  • Into the Jungle: Great Adventures in the Search for Evolution (2008, Benjamin Cummings; ISBN 0-321-55671-2)
  • Remarkable Creatures: Epic Adventures in the Search for the Origin of Species (2009, Houghton Mifflin Harcourt; ISBN 0-15-101485-X)
  • Brave Genius: A Scientist, a Philosopher, and Their Daring Adventures from the French Resistance to the Nobel Prize (2013, Crown; ISBN 0-307-95233-9)
  • The Serengeti Rules: The Quest to Discover How Life Works and Why It Matters (2016, Princeton University Press, ISBN 9780691167428)

Magazine articles

  • The Origins of Form: Ancient genes, recycled and re-purposed, control embryonic development in organisms of striking diversity (2005, Natural History Magazine) [18]
  • God as Genetic Engineer. A review of Michael Behe's book "The Edge of Evolution: The Search for the Limits of Darwinism" (2007, Science Magazine) [19]
  • Regulating Evolution: How Gene Switches Make Life (2008, Scientific American) [20]


Science writer Peter Forbes, writing in The Guardian, calls Endless Forms Most Beautiful an "essential book" and its author "both a distinguished scientist ... and one of our great science writers." In Forbes's view, in The Serengeti Rules Carroll "manages to unite natural history with the hard science of genomics."[21] In her article on Science Based Medicine titled The Essential Role of Regulation In Human Health and In Ecology: The Serengeti Rules, Harriet Hall says "This book is a great way to learn about the rules of regulation and about how science works. It’s not just a painless way to learn, it’s positively fun."[22] The documentary film, The Serengeti Rules, was released in 2018 and is based on Carroll's book.[23]

Louise S. Mead, reviewing The Making of the Fittest for the National Center for Science Education, notes that Carroll provides "some of the overwhelming evidence for evolution provided in DNA", using different lines of inquiry such as DNA sequences that code for genes no longer in use, and evidence of evolutionary change. Mead notes that evolutionary theory has predictive power, as with icefish whose ancestors had haemoglobin, but no longer needing it in icy water, they have lost it.[24]

Douglas H. Erwin, reviewing Endless Forms Most Beautiful for Artificial Life, remarks that life forms from Drosophila to man have far fewer genes than many biologists expected – in man's case, only some 20,000, which is about the same as a fly. He notes the "astonishing morphological diversity" of animals coming from "such a limited number of genes". He praises Carroll's "insightful and enthusiastic" style, writing in a "witty and engaging" way, pulling the reader into the complexities of Hox and PAX-6, as well as celebrating the Cambrian explosion of life forms, and much else.[25]


  2. "Benjamin Franklin Medal in Life Science". Franklin Institute. 2012. Retrieved April 6, 2013.
  3. "Our Scientists". HHMI. Howard Hughes Medical Institute. Retrieved August 11, 2017.
  4. "LCMB Investigators". Laboratory of Cell and Molecular Biology at UW-Madison. Retrieved May 2, 2017.
  5. Wade, Nicholas (July 5, 1994). "How Nature Makes a Butterfly's Wing". The New York Times. p. C9. Retrieved May 4, 2017.
  6. "Shaw Scientist Award Recipients". Greater Milwaukee Foundation.
  7. "NOVA: Family That Walks on All Fours". PBS.
  8. Carroll, Sean B. "In a Shark's Tooth, a New Family Tree". The New York Times. Retrieved February 10, 2017.
  9. Carroll, Sean B. "Solving the Puzzles of Mimicry in Nature". The New York Times. Retrieved February 10, 2017.
  10. "Sean B. Carroll, HHMI Vice President for Science Education". Howard Hughes Medical Institute. Retrieved October 22, 2013.
  11. "HHMI Launches Documentary Film Unit to Create Science Features for Television". HHMI. Retrieved May 2, 2017.
  12. "HHMI Premieres New Film Showcasing One of Science's Greatest Detective Stories". HHMI. Retrieved May 2, 2017.
  13. "The Stephen Jay Gould Prize". Society for the Study of Evolution. Retrieved May 2, 2017.
  14. "SSE 2010 Stephen Jay Gould Prize".
  15. "Presentation of the 2016 Lewis Thomas Prize to Sean B. Carroll". The Rockefeller University. Retrieved April 15, 2016.
  16. Carroll, Sean B (2008). "Evo-Devo and an Expanding Evolutionary Synthesis: A Genetic Theory of Morphological Evolution". Cell. 134: 25–36. doi:10.1016/j.cell.2008.06.030. PMID 18614008.
  17. "Advisory Council". National Center for Science Education. Archived from the original on August 10, 2013. Retrieved October 30, 2018.
  18. Carroll, Sean B. "The Origins of Form". Natural History Magazine. Retrieved May 2, 2017.
  19. Carroll, Sean B (June 8, 2007). "God as Genetic Engineer". Science Magazine. 316 (5830): 1427–1428. doi:10.1126/science.1145104.
  20. Sean B Carroll; Nicolas Gompel; Benjamin Prudhomme (May 2008). "Regulating Evolution: How Gene Switches Make Life". Scientific American. Retrieved May 3, 2017. (Preview)
  21. Forbes, Peter (March 23, 2016). "The Serengeti Rules by Sean B Carroll review – a visionary book about how life works". The Guardian. Retrieved April 15, 2016.
  22. Hall, Harriet. "The Essential Role of Regulation In Human Health and In Ecology: The Serengeti Rules". Science Based Medicine. Retrieved March 1, 2016.
  23. Regan, Chelsea (April 7, 2019). "PBS International & HHMI Tangled Bank Partner on Science Docs". TVREAL. Retrieved April 23, 2019.
  24. Mead, Louise S. (2008). "Review: The Making of the Fittest". Reports of the National Center for Science Education. 28 (1): 37–39.
  25. Erwin, Douglas J. (2007). "Book Review: Endless Forms Most Beautiful". Artificial Life. 13 (1): 87–89.
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