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Highlights of the research careers of John and Elisabeth Buck are presented, illustrating their importance as investigators of firefly biology including taxonomy, morphology, physiology and behavior, and as catalysts of collegial exchanges advancing progress among investigators of bioluminescence in its widest aspects over the past 50 years.
The mysterious process by which fireflies can control their flashing has inspired over a century of careful observation but has remained elusive. Many studies have implicated oxygen as the controlling element in the photochemical reaction, and the discovery of nitric oxide synthetase (NOS) in the lantern has suggested that nitric oxide (NO) may control oxygen access to the light-emitting photocytes, thereby triggering the flash. However, there are several drawbacks to oxygen as a controlling agent, and in view of the prominence of peroxisomes in lantern morphology and biochemistry, we suggest that it is hydrogen peroxide that triggers the flash, and we present a model by which this may take place.
In light-producing cells (photocytes) of the firefly light organ, mitochondria are clustered in the cell periphery, positioned between the tracheolar air supply and the oxygen-requiring bioluminescent reactants which are sequestered in more centrally-localized peroxisomes. This relative positioning suggests that mitochondria could control oxygen availability for the light reaction. We hypothesized that active cellular respiration would make the interior regions of the photocytes relatively hypoxic, and that the “on” signal for production of bioluminescence might depend on inhibition of mitochondrial oxygen consumption, which would allow delivered oxygen to pass through the peripheral mitochondrial zone to reach peroxisomes deep in the cell interior. We published recently that exogenous NO induces bioluminescence in the intact firefly; that NO mediates octopamine-induced bioluminescence in the dissected lantern, and that nitric oxide synthase is abundant in cells of the tracheolar system of the light organ. Additional experiments showed that nitric oxide gas (NO) inhibits respiration in isolated lantern mitochondria. Inhibition is reversed by bright light, and this inhibition is relieved when the light is turned off. Altogether, the results support the idea that NO triggers light production by reversible inhibition of mitochondrial respiration in lantern cells, and probably in tracheolar cells as well. The data also suggest that the light of bioluminescence itself relieves NO inhibition thus contributing to rapid on/off switching. While other mechanisms may be in play, NO production that is directly related to neural input appears to have a key role in the oxygen gating that controls flash communication signals.
The very different courtship flashes of Photuris versicolor and Photuris lucicrescens males mirror the pattern of neural impulses produced by their brain. Their lanterns luminescence very differently, however, in response to direct, electrical stimulation. Whereas P. lucicrescens lanterns glow in response to high frequency, continuous electrical stimulation, those of P. versicolor produce only rapid, triple-pulsed flashlettes that resemble, but are not identical to, their courtship flashes. In addition, the exposed lantern tissue of P. versicolor males, when immersed in firefly saline high in potassium and calcium ions, scintillates with hundreds of photocytes flashing in random fashion. P. lucicrescens male lanterns, so treated, only glow. Tests of P. versicolor lanterns with salines of different composition suggest that calcium ions are essential in producing this intense, long lasting scintillation response and are therefore possibly implicated in the final stages of flash control in this species.
Japanese fireflies range from nocturnal luminescent species to diurnal non-luminescent species. Their communication systems are classified into 6 types based on the following criteria: 1) Female responds to male's flashes after a fixed delay; 2) Male is directly attracted by female's light signal, the male perches on a leaf near the female, then the male changes his flashes with twinkling, and copulation behavior is released. However, the female may not respond to the male; 3) Male seeks female calling signal during the male's flying and synchronous flashing, then the male approaches the female, emitting flashes with various patterns, displaying walking-luminescing, sedentary signaling, chasing, and copulating; 4) Male is attracted by continuous luminescent signals of the female, and male perches near the female, then the male distinguishes the female's light organs shape. Thereafter, the male copulation behavior is released by her sex pheromone; 5) Male and female flight occurs in the daytime; when the male approaches the female, copulation is released by the female's pheromone; weak luminescent signals may be fulfilling the function of supplementary communication signals; 6) Luminescent signals have nothing to do with communication between male and female, and copulation is released by a sex pheromone.
The phenomenon of nuptial gift transfer during mating occurs across a remarkably wide range of taxa, and such male donations are likely to influence both pre-copulatory and post-copulatory sexual selection. This paper reviews what is known about nuptial gifts in Photinus fireflies (Coleoptera: Lampyridae), and discusses the adaptive significance of spermatophores in firefly mating systems. During copulation Photinus males transfer a spiral, gelatinous spermatophore to the female: sperm are released into the female's spermatheca for storage, while the remainder of the spermatophore disintegrates within a specialized gland. Radiolabelling studies indicate that male-derived protein is used to help provision the female's developing oocytes, and multiply-mated females show increased fecundity. As most Photinus adults do not feed, these studies suggest that females should continue to forage for matings to supplement their diminishing larval reserves, even after they have gained sufficient sperm to fertilize their eggs. Male spermatophore mass declines across sequential matings, and smaller spermatophores are associated with lower paternity success in situations where males compete for fertilizations. Declining spermatophore size across sequential matings may thus lead to diminishing reproductive returns for firefly males. Taken together, these results suggest that seasonal changes in nuptial gift availability may contribute to reversals of traditional courtship roles, with male choice and female-female competition occurring as spermatophore availability declines.
The evolution of male courtship signals such as the bioluminescent flashes of fireflies may be shaped, at least in part, by female preference for particular characteristics of the male signal. These female preferences for male courtship signals may arise as a result of the benefits of choosing males with particular traits. One possible benefit of mate choice occurs if females can use male courtship signals as an honest indicator of male nutritional contributions at mating, nuptial gifts. This paper reviews female preference for male flash characteristics in Photinus fireflies (Coleoptera: Lampyridae), and the potential for females to use male flash characteristics to predict nuptial gift quality. In Photinus firefly species with single pulse flashes females preferentially respond to flashes of greater intensity and duration. Male Photinus provide a nuptial gift to females at mating in the form of a spermatophore and flash duration serves as a good predictor of spermatophore mass for males collected early in the season. However, Photinus fireflies do not feed as adults, so spermatophore mass decreases with subsequent matings. In response, nutrient-limited females may stop preferentially responding to longer duration flashes, increasing their overall responsiveness later in the mating season as they forage for spermatophores. Therefore, the evolution of male courtship signals in Photinus fireflies is the product not only of female preference for male flash characteristics, but also the costs and benefits of female choice that shape these preferences.
Ever since Darwin identified it as the force responsible for the evolution of exaggerated male characters, sexual selection has been the focus of research aimed at understanding the most bizarre and intriguing morphologies and behaviors in Nature. Two congeneric species in the firefly genus Photinus, P. pyralis and P. macdermotti, afford a unique opportunity to examine the interaction between sexual and countervailing natural selection that act to shape the evolution of mating behavior and body size in closely related species with very different courtship strategies. Photinus pyralis males emit very bright flashes during their extended patrolling flights and form ‘love knots’ of competing suitors, while P. macdermotti males, whose courtships seldom exceed two competing males, produce weaker flashes during a shorter patrolling period.
Possibly as a consequence of their scramble competition and long flights, not only is there an extremely wide range of body sizes in P. pyralis males, but they also exhibit wing allometric slopes greater than one, and lantern allometries less than one. In contrast, P. macdermotti males do not have allometric slopes significantly different from one. Small males, when tested in an artificial scramble situation move faster than large males, an advantage in the intense competition that ensues once a female is located. Females answer several males in alternation and fail to respond to all male flashes, with the evident consequence that love knots frequently develop around such fickle females. Allometric relationships in two non-luminescent beetle species with non-visual courtship protocols are compared. Visual, sexually selected characters showed positive allometric slopes, while non-visual characters showed isometry or negative slopes. Data presented here support the existence of distinct patterns in modality-specific sexual selection.
Flash communication by the firefly Photinus pyralis was studied in a stationary, simulated flight apparatus in which an individual of either sex could be “flown” and its flashing behavior and flight orientation recorded in response to photic stimulation. Males made long “flights” showing many of the characteristics of their natural, female-seeking patrol flights. Males oriented their flight vectors towards light emitting diode (LED) flashes that mimicked the responses of females to their patrol flashes. Females flew and responded to male-emulating LED flashes, making a previously unknown early response followed by the typical 2 sec delayed response characteristic of the dialoging perched female, including abdominal aiming of the flash. Pairs consisting of males, in tethered flight, and females, perched, were run in an integrating sphere photometer, permitting the first determinations of flash intensities of both sexes during courtship dialog. The implications of this work on thought about evolution of photic behavior in fireflies are considered.
Synchronous flashing occurs in certain species of Southeast Asian and North American fireflies. Most Southeast Asian synchrony involves stationary congregating fireflies, but North American synchrony occurs in flying fireflies that do not congregate. Southeast Asian synchrony is usually continuous, but North American synchrony is interrupted. Photuris frontalis, the only member of the North American genus Photuris to synchronize, shows an intermittent synchrony. This involves synchronization and repeated re-synchronizations while in flight. The precision that occurs at the start of synchrony was studied in Ph. frontalis using caged fireflies and photometry. Barrier experiments (using two fireflies) or flash entrainment experiments (using one LED and one firefly) were performed to measure the temporal precision of the first entrained flash. In both cases, the first entrained flash was close to unison synchrony (phase = 1.0) and showed little variability. The behavioral implications of the ability to synchronize with the first entrained flash are not known, but it might facilitate male-male interactions during brief, transient encounters such as maintaining distance between closely flying males in search of females.
The study of fireflies makes an ideal subject for introducing students of all ages to the world of science, conservation biology, field studies and the importance of observation and understanding connections in the natural world.
The synchronous fireflies, Photinus carolinus of the Great Smoky Mountains National Park's Elkmont Historic District can be used as a basis for classroom and field activities in subjects as diverse as conservation biology, economics, geography, ecology, governmental policy, zoology, history, taxonomy and animal behavior.
Having been fascinated with the synchronous fireflies for over two decades, I will use the life cycle of the Elkmont Phontinus carolinus found in the Great Smoky Mountains National Park as an example.