User:Turnbull.41/sandbox

For my topic I will be exploring the evolution of light patterns of fireflies with natural selection and mating.

Lewis, Sara M. Cratsley, Christopher K.  2008. Flash Signal Evolution, Mate choice, and Predation in Fireflies. Annual Review of Entomology. 53: 293-321. Available from: http://ase.tufts.edu.proxy.lib.ohio-state.edu/biology/labs/lewis/publications/documents/2008Lewis_Cratsley.pdf

-This article deals with the mate choice of fireflies according to their light pattern along with the evolution of different light patterns that differentiate species from one another, a form of behavioral isolation.

Biggley, W. H., Lloyd, J. E., Seliger, H. H. 1967. The Spectral Distrobution of Firefly Light II. JGP. 50(6): 1681-1692. Available from: http://jgp.rupress.org.proxy.lib.ohio-state.edu/content/50/6/1681.full.pdf+html

-This article deals with the spectrum of light that fireflies deal with with accordance to what the color of the organs are in their bodies, along with luciferase molecules and the role that they play in the light pattern and light colors of fireflies.

Lloyd, James E. 1971. Bioluminescent Communication in Insects 6004. Annual Review of Entomology. 16: 97-122. Available from: http://www.annualreviews.org.proxy.lib.ohio-state.edu/doi/pdf/10.1146/annurev.en.16.010171.000525

-This article talks about the communication between insects, including fireflies, because of bioluminescence and that the light pattern and color are not only dependent on mate choice between species, but also the communication between species and in the species.

Orlova, Galina., Goddard, John D., Brovko, Lioubov Yu. 2003. Theoretical Study of the Amazing Firefly Bioluminescence: The Formation and Structures of the Light Emitters. J. Am. Chem. Soc. 125(23): 6962-6971. Available from: http://pubs.acs.org.proxy.lib.ohio-state.edu/doi/full/10.1021/ja021255a

-This article seems useful because it talks about how the light is emitted and transformed in fireflies. This can be a big breakthrough on how different species arise in fireflies along with their behavior, mate choice, and the chemistry/biology aspect of how these light emitters are structured.

Eisner, Thomas., Goetz, Michael A., Hill, David E., Smedley, Scott R., Meinwald, Jerrold. 1997. Firefly "feemes fatales" acquire defensive steroids (lucibufagins) from their firefly prey. 94(18): 9723-9728. Available from: http://www.pnas.org.proxy.lib.ohio-state.edu/content/94/18/9723.full

-This article could be helpful because it talks about how females of one species of fireflies mimic an action of another species, while trapping males of the other species in this trap so they can become the females prey. Also the females gain a steroid called the lucibufagin which helps them against the predators of jumping spiders which reject this steroid, so this can play a huge role in natural selection.

Project Part 2
1. https://en.wikipedia.org/wiki/Talk:Bioluminescence 2. https://en.wikipedia.org/wiki/Talk:Mating 3. https://en.wikipedia.org/wiki/Talk:Firefly

This was my actual edit to the Bioluminescence page. Don't have nearly as enough information as I wanted there yet, but it will get there. Also I'm not quite sure if I did the citation right on the page? https://en.wikipedia.org/wiki/Bioluminescence#Mimicry

Final Draft Starts Here!
Fireflies are a very interesting species when they come into the conversation of evolutional biology. There are many aspects of fireflies that make them very unique and intriguing to other species such as the light patterns they form at night and how this is dependent on differentiating each other from different species and for mating purposes. Along with this, every species has its consequences with the environment such as mimicking from other species. What will be discussed is the effect of behavioral isolation in the differences in light pattern from species to species, the spectrum of light fireflies emit depending on the color of their organs and the abundance of luciferase molecules, how light pattern is not only dependent on mate choice but also communication between a species or others, how different fireflies with different light patterns evolved or diverged from a common ancestor and the structure of their light emitters, and how females mimic the effects of other species in order to trap their prey the males and how they gain a steroid called lucibufagin to help protect themselves against predators. Both male and female fireflies produce flash signals in order to communicate with one another for courtship purposes (Lewis and Cratsley 2008). The communication usually starts with the male while it is flying making signals at the same time to grab the attention of females. When the females respond this usually ends up in courtship of the males and females and they copulate. However, there are different flash timings and signals according to different species of fireflies, but perhaps the idea is that males start with advertising flash signals. When females are attracted to that signal they respond to the male where the male is attracted to their response signal, they both communicate through signals and timing and it all ends in copulation if they are successful (Lewis and Cratsley 2008). For North American fireflies there are three different types of modes that they use for signaling which are diurnal fireflies use pheromones during flight in the daytime, glowworm male flies look for female flies which are in larva form and be seen by their bioluminescence in burrows that attract the males. There are also flying males that communicate with females at nighttime due to signals ending up in copulation (Choe and Crespi 1997). The differences in these mating types along with different signaling techniques could provide us with some of the evolutionary history fireflies have gone through in accordance to selective pressures (Choe and Crespi 1997). In glowworms, for example, the males don’t emit the light, but the females do. Along with any species, but perhaps glowworms in general this could lead to more male-male competition due to the females emitting light in the burrow. As for the diurnal fireflies they search during the daytime, which might explain why they do not have bioluminescence because the light would not be able to be seen during the daytime. However, they do mate with the females at night while the females are buried underground, but perhaps this is where the bioluminescence would be “preselected” in order to notice the light or flash signals (Choe and Crespi 1997). Sometimes the size, shape, pattern, and timing of the communication occuring between individual organisms is not the only factor that plays into bioluminescent communication. Movement can play a role between fireflies, but it is not significant to the mating process. Sometimes while males and females are trying to communicate to each other with regards to successful copulation, the male makes species specific movements in order to mate with the female. Studies show that this is perhaps now true according to species, however it could be significant in regards to other insects who may or may not be bioluminescent like fireflies (Lloyd 1971). Depending on the region where each species is typically inhabited or what time during the night it currently is can have an effect on the signals fireflies make according to color. It was observed from North American fireflies the ones that are more active while it is dark out, or in the middle of the night, emit green light while the fireflies more active while at dusk emit yellow light (Viviani and Bechara 1995). Along with the fireflies that are more active at dusk, it is said that they emit the yellow light in order to compensate for the green color the grass gives off, but isnot yet fully understood (Viviani and Bechara 1995). While comparing the Brazilian species to the North American species it was found that the Brazilian species were more abundant in yellow-green and green emission as North American species prevails in yellow emission (Viviani and Bechara 1995). This shows a clear difference in the location of species and their difference in possibly light patterns, but most definitely on the color of their bioluminescence, which could be an example of allopatric speciation. A study was conducted to obtain wavelengths of light and color according to different species such as American fireflies and Jamaican fireflies. The study was conducted by using a small sharp needle and inserting it into the firefly’s abdomen where the light emissions were coming from in order to look at the organs and get a better look at their organ processes. It was found that between Jamaican and American fireflies that they do indeed form quite different wavelengths of light and color while Jamaican fireflies are more prone to lower wavelengths which are a little darker in color and the American fireflies were more likely to have higher wavelengths, which is brighter colors (Biggley et al. 1967). These differences between the Brazilian and North American fireflies along with Jamaican and American fireflies are a great example and huge possibility that this could be evidence of allopatric speciation, or in other words, geographic isolation. This means that these species at one point in time were probably part of the same population when the landmasses were more connected to each other. Now since North America and South America along with America and Jamaica are separated by large bodies of water, there is a good explanation why these two species went through an allopatric speciation event. One of the main selective pressures against fireflies is predation because of their greater possibility to be seen more often due to their flash signals and the emission of light (Magnhagen 1991). However, some fireflies had made adaptions to this predation by instead of going through copulation multiple times, to make the process quicker but possibly less ineffective is to copulate only one at a time to decrease the occurrence of predation (Magnhagen 1991). There is also evidence of an arms race with fireflies and their predators, for example there are some species of firefly predators that mimic the effect of a certain signaling method to lure in their prey. What it means for the fireflies and their predators to coevolve in this way and example, the approaching male who is falling for the predators mimicry can perhaps shut off it’s emission of light completely (Magnhagen 1991). Along with the predation of fireflies there is one in particular of a male species that when hunting for males gain a specific steroid to help protect itself from jumping spiders. Females of the Photuris species were found to contain the steroid of lucibufagin (LBG), but it was found that they actually gain this defensive steroid from males of another firefly species, Photinus, that naturally produce this steroid. Photuris females lure in this different species of males by mimicking the effect of their light-signaling pattern. The males are fooled because they think this is the response of a female from their same species. This eventually ends up having the Photinus male being eaten by the Photuris female, in turn giving the female the steroid lucibufagin (Eisner et al. 1997). So how does this help with the females defense against jumping spiders? A study was performed where the Photuris females were collected from nature and forced to reflex bleed which contains the steroid lucibufagin. It was found that when the females were forced to reflex bleed, the samples taken from each female had different amounts of the steroid in each sample. So after experiments were brought out to see which females the jumping spiders would eat it was decided that the jumping spiders were more likely to eat the females with less lucibufagin inside their bodies and the females with more were constantly rejected by the spiders therefore protecting themselves from predation (Eisner et al. 1997). There is a lot more information on how fireflies emit light from their bodies. There is actually some chemistry involved with the emission of light. For bioluminescent and fireflies alike the bioluminescence actually comes from the oxidation of a substrate inside the insect by an enzyme. The substrate is luciferin, LH2, which is then catalyzed by the enzyme of luciferase. This enzyme in turn makes the molecules of luciferin in an excited state that makes the bioluminescent signals of some insects including fireflies (Orlova et al. 2003). Since the bioluminescence in insects such as fireflies deals with the production of light in their bodies there has to be some sort of energy input and output due to this light emission. ATP (adenosine triphosphate) is naturally occurring energy in nature and perhaps helps with the production of light in fireflies. In this specific example ATP is the energy transferring source in bioluminescent insects and actually increases the reaction of the oxidation of luciferin most likely causing the light emitted from fireflies to be so visible (Orlova et al. 2003). To summarize, fireflies have been through a lot of evolutionary forces such as behavioral isolation due to differences in the shape, timing, pattern, and movement of the light that they emit. Along with this behavioral isolation can be due to the fact of how males and females of the same species interact with each other on how they communicate due to flash signals, and then eventually copulate when successful with the communication. Also, there is a chance that due to evolutionary differences between fireflies of North America and Brazil, these fireflies went through an allopatric speciation event because of their differences in wavelengths and colors they emit. Between the fireflies and their predators there is a constant coevolution between them, or an arms race where the fireflies are constantly adapting to the predators new strategies such as changes in the amount of time males spend copulating with females as to not waste time. References Biggely, W. H. Lloyd, J. E.  Seliger, H. H.  1967. The Spectral Distribution of Firefly Light II. JPG [Internet]. 50(6): 1681-1692. Available from: http://jgp.rupress.org.proxy.lib.ohio-state.edu/content/50/6/1681.full.pdf+html Choe, Jae C. Crespi, Bernard J.  1997. Firefly Mating Ecology, Selection and Evolution. Mat Syst in Ins and Arach [Book from Internet]. Cambridge (UK). Cambridge University Press. Available from: http://books.google.com.proxy.lib.ohio-state.edu/books?hl=en&lr=&id=X3Iz-zXolVcC&oi=fnd&pg=PA184&dq=Firefly+evolution&ots=yNoCHNnrKK&sig=q8LDEPfYztYUpPiQVscKrlRAQU0#v=onepage&q=Firefly%20evolution&f=false Eisner, Thomas. Goetz, Michael A. Hill, David E.  Smedley, Scott R.  Meinwald, Jerrold. 1997. Firefly “feemes fatales” acquire defensive steroids (lucibufagins) from their firefly prey [Internet]. 94(18): 9723-9728. Available from: http://www.pnas.org.proxy.lib.ohio-state.edu/content/94/18/9723.full Lewis, Sara M. Cratsley, Christopher. 2008. Mate Choice and Predation in Fireflies. Annul Rev Etom [Internet]. 53: 293-321. Available from: http://ase.tufts.edu.proxy.lib.ohio-state.edu/biology/labs/lewis/publications/documents/2008Lewis_Cratsley.pdf Lloyd, James E. 1971. Bioluminescent Communication in Insects 6004. Annul Rev Etom [Internet]. 16: 97-122. Available from: http://www.annualreviews.org.proxy.lib.ohio-state.edu/doi/pdf/10.1146/annurev.en.16.010171.000525 Magnhagen, Carin. 1991. Predation Risk as a Cost of Reproduction. Tren In Ecol and Evol [Internet]. 6(6): 183-186. Available from: http://ac.els-cdn.com.proxy.lib.ohio-state.edu/016953479190210O/1-s2.0-016953479190210O-main.pdf?_tid=4aeab848-5f16-11e4-ae11-00000aacb361&acdnat=1414551187_6dc234824b3a48560090a36850e966b4 Orlova, Galina. Goddard, John D. Brovko, Lioubov Yu. 2003. Theoretical Study of the Amazing Firefly Bioluminescence: The Formation and Structures of the Light Emitters. Jour of Amer Chem Soc [Internet]. 125(23): 6962-6971. Available from: http://pubs.acs.org.proxy.lib.ohio-state.edu/doi/full/10.1021/ja021255a Viviani, Vadim R. Bechara, Etelvino J. H.  1995. Bioluminescence of Brazilian Fireflies (Coleoptera: Lampyridae): Spectral Distribution and pH Effect on Luciferase-Elicited Colors. Comparison with Elaterid and Phengodid Luciferases. Photochem and Photobio [Internet]. 62(3): 490-495. Available from: http://onlinelibrary.wiley.com.proxy.lib.ohio-state.edu/store/10.1111/j.1751-1097.1995.tb02373.x/asset/j.1751-1097.1995.tb02373.x.pdf?v=1&t=i1u3cshd&s=6f01f6645b321fc970fe209688eecc38a9b71cad

Final Page Edits
1. https://en.wikipedia.org/wiki/Photinus_pyralis The second paragraph in the "Defense" section.