Notes 6: The Making of a Flasher

A Chemistry Set for Light
Reviews by Wilson & Hastings (1998, 2013) outline the chemistry of bioluminescence. The rendition of luciferase is from David S. Goodsell’s June 2006 Molecule of the Month in the RCSB’s Protein Data Bank (

Firefly Lights Evolving
Viviani (2002) and Oba (2015) review ideas about how beetle luciferases might have evolved. Yuichi Oba and his colleagues (2008) measured luciferin content of various non-luminescent beetles. Lynch (2007) describes the role of gene duplication in the evolution of snake venom. Darwin’s prescient quote about exaptation is from the Origin of Species (1859, p.190).

Putting Fireflies to Work
Practical applications for fireflies’ light-producing ability are described in Weiss (1994), Rosellini (2012) and Andreu and colleagues (2013).

Weiss, R. (1994, August 29) Researchers gaze into the (insect) light and gain answers, The Washington Post: A3.

Controlling the Flash
John Buck’s biographical information is from Case & Hanson (2004), and from his New York Times obituary.

Pearce, J. (2005, April 3). John B. Buck, who studied fireflies’ glow, is dead at 92. The New York Times.

A Journey Inside Firefly Lantern
Buck (1948) and Ghiradella (1998) provide a detailed look at the internal anatomy of the firefly lantern. Helen Ghiradella is not only an expert on firefly lantern anatomy, but also a skilled artist whose drawings allow others to appreciate the internal architecture of this light-producing marvel. Figure 6-2 is modified with permission from Ghiradella (1998).

Discovering the Firefly’s Light Switch
Our discoveries about the role of nitric oxide in firefly flash control were reported in Trimmer et al. (2001). A complementary hypothesis about oxygen control based on the anatomy of firefly airtubes is presented by Ghiradella & Schmidt (2008).

Getting in Sync
Smith (1935) describes witnessing the synchronous fireflies (Pteroptyx) in Thailand, although he incorrectly believed this activity was unrelated to mating.

John and Elisabeth Buck’s trip to Thailand produced the very first scientific study of how Pteroptyx malaccae fireflies manage to flash synchronously (Buck & Buck 1968). Elisabeth Buck’s quote is from the Radiolab podcast, Emergence broadcast on February 18, 2005. Bookending 50 years of scientific research on the mechanisms of firefly flash synchrony, John Buck wrote two review articles about the different physiological mechanisms underlying flash synchrony: Buck (1938) and Buck (1988). In the latter, he also describes the geographic and taxonomic distribution of several types of flash synchrony.

Mathematician Steve Strogatz’s book, Sync: The Emerging Science of Spontaneous Order, provides an entertaining and accessible description of how pulse-coupled oscillators lead to synchrony in the engineered and natural world (Strogatz 2003).

Science Confidential
Niko Tinbergen (1907 – 1988) was a Dutch ethologist and ornithologist who shared the 1973 Nobel Prize in Physiology or Medicine for his discoveries about animal behavior. His 1963 paper was dedicated to Konrad Lorenz on the ocassion of his 60th birthday. My account of this scientific feud is based on personal conversations and correspondence exchanged with both Jim Lloyd and John Buck. Buck & Buck (1978) center their explanations on how synchrony could benefit a group of males, while Lloyd (1973b) presents some ideas about what advantages flash synchronization could provide both to individual males as well as to the group. Faust (2010) describes how P. carolinus males switch from synchronizing their flashes to chaotic flashing once they approach a female. Case (1980) describes some close-up behavioral interactions inside Pteroptyx display trees. 

Further Exploration

More about Bioluminescence
Living creatures that make their own light are deeply fascinating. The 2009 science fiction movie Avatar, written and directed by James Cameron, is well-known for the exuberantly bioluminescent flora and fauna that inhabit the fantasy world called Pandora. Two leading authorities on bioluminescence, Thérèse Wilson and Woody Hastings, describe the molecular details of bioluminescence in selected creatures, including fireflies.

Wilson, T. and J. W. Hastings (2013). Bioluminescence: Living Lights, Lights for Living. Harvard University Press, Cambridge MA. 208 pp.

The Bioluminescence Web Page celebrates all living things that produce their own light (hosted by the University of California at Santa Barbara).

More about Synchrony
Writing for a general audience, leading mathematician and an award-winning science communicator Steve Strogatz explains how thousands of fireflies, cardiac pacemaker cells, or electrons in a superconductor manage achieve their highly organized synchronous behavior without a conductor.

Strogatz, S.H. (2003). Sync: The Emerging Science of Spontaneous Order. Hyperion Books, NY. 338 pp.

Biologist Michael Greenfield takes a detailed look at the acoustic, chemical, and vibratory, visual, and bioluminescent signals used by insects. He also reviews what we know about the hows and whys of collective male synchrony in fireflies, crickets, and cicadas.

Greenfield, M. (2002). Signalers and Receivers: Mechanisms and Evolution of Arthropod Communication. Oxford University Press, NY. 432 pp.

Radiolab: Emergence

This February 18, 2005 Radiolab podcast explores how individuals following simple rules can generate complex group behaviors, like firefly synchrony. This episode features interviews with biologists John and Elisabeth Buck, as well as with mathematician Steve Strogatz.