User:Leahyaa/Slendertail lanternshark/Bibliography

Bibliography - Amanda

 * Claes, Julien M., Nilsson; Dan-Eric.; Straube, Nicolas. (2014). "Iso-luminance counterillumination drove bioluminescent shark radiation". Sci Rep.
 * details on the photophores of E. molleri
 * Duchatelet, L., Pinte, N., & Tomita, T. (2019). Etmopteridae bioluminescence: dorsal pattern specificity and aposematic use. Zoological Letters.
 * pattern of photophores on E. molleri's body
 * Duchatelet, L., Claes, J. M., & Delroisse, J. (2021). Glow on Sharks: State of the Art on Bioluminescence Research. Oceans.
 * Details on photophore arrangement and regions created on the body plan by said patterns.
 * Mizuno, G., Yano, D., & Paitio, J. (2021). Etmopterus lantern sharks use coelenterazine as the substrate for their luciferin-luciferase bioluminescence system. Biochemical and Biophysical Research Communications.
 * details on bioluminescence of slendertail sharks being due to Coelenterazine activity
 * Claes, J. M., Mallefet, J. (2015). Comparative control of luminescence in sharks: New insights from the slendertail lanternshark (Etmopterus molleri). Journal of Experimental Marine Biology and Ecology.
 * general information of photophore activity in E. molleri and patterning on slendertail shark

Bibliography - Brendan

 * Duchatelet, L., Pinte, N., Tomita, T., Sato, K., Mallefet, J. (2019). Etmopteridae bioluminescence: dorsal pattern specificity and aposematic use. Zoological Letters, pp. 1-10.
 * Dorsal luminescence in E. molleri is useful for counterillumination, possibly as a defense mechanism.
 * This paper suggests that bioluminescence is aposematic, fighting against predator attractiveness and hunting.
 * Other species were discussed, including the second species of Etmopterus that our group is studying. This could suggest behavioral similarities between lantern sharks that have evolved similar styles of bioluminescence.
 * Claes, J., Mallefet, J. (2015). Comparative control of luminescence in sharks: New insights from the slendertail lanternshark (Etmopterus molleri). pp. 87-94.
 * This paper discusses bioluminescence in E. molleri as beneficial for camouflage in addition to bioluminescent signaling, as well as relating to other species who could use bioluminescence for sexual signaling. This is important, and I’d like to explore more about reproductive strategies of E. molleri, and if bioluminescence is involved at all.
 * However, this paper also does address the fact that ecology of E. molleri is mostly undiscovered otherwise, which means there may be no relationship between bioluminescence and sexual signaling.
 * Duchatelet, L., Claes, J., Delroisse, J., Flammang, P., Mallefet, J. (2021). Glow on Sharks: State of the Art on Bioluminescence Research. MDPI - oceans. pp. 822-842.
 * This source acts as another bolster of support to the aposematic theory. E. molleri presents bioluminescence on spines atop of dorsal fins. However, these spines also double as physical defense against predators aside from bioluminescence.
 * Chemical evidence shows that bioluminescence in E. molleri is triggered, not constitutively active. This proves, though not completely clear, that behavioral stimuli must be met for bioluminescence to occur.
 * Mallafet, J., Stevens, D., Duchatelet, L. (2021). Bioluminescence of the Largest Luminous Vertebrate, the Kitefin Shark, Dalatias licha: First Insights and Comparative Aspects. Frontiers in Marine Science.
 * In a Kitefin Shark, the pelvic area is bioluminescent, but not bright enough to make an impact. In E. molleri, bioluminescence outlights the rest of the body, and this article assumes that this is more important for sexual signaling than the Kitefin shark’s bioluminescence is.
 * This article also provides supporting evidence towards the aposematic benefits of bioluminesce in E. molleri, as well as counterillumination.
 * Duchatelet, L., Sugihara, T., Delroisse, J. Koyanagi, M., Rezsohazy, R., Terakita, A., Mallefet, J. From extraocular photoreception to pigment movement regulation: a new control mechanism of the lanternshark luminescence. Nature.com Scientific Reports. pp. 1-15.
 * This article describes ventral bioluminescence from E. molleri, mostly as a camouflage mechanism.
 * Other benthopelagic lanternsharks also express bioluminescence in a similar manner, suggesting that the mechanism by which this occurs may have evolved together.

Bibliography - Dayna
Claes, J. M., Krönström, J., Holmgren, S., & Mallefet, J. (2010). Nitric oxide in the control of luminescence from Lantern Shark (Etmopterus Spinax) photophores. Journal of Experimental Biology, 213(17), 3005–3011. https://doi.org/10.1242/jeb.040410


 * The authors found evidence that nitric oxide (NO) moderates the expression of luminescence in photocytes by acting on hormones. They collected specimens of Etmopterus spinax and analyzed their photophores for reactivity to NO through testing whether injecting nitrergic drugs would produce luminescence by itself, and then administered hormones involved in the production of light including melatonin (MT) and prolactin (PRL) to observe the effect. Alone, the nitrergic drugs did not produce luminescence but in conjunction with MT and PRL altered the production of luminescence. Although this study was conducted with E. Spinax and not Etmopterus molleri, this study discovered the mechanism by which NO moderates hormone expression of luminescence which has also been replicated in an experiment with E. Molleri that I am also using in my research (Comparative control of luminescence in sharks: New insights from the slendertail lanternshark (Etmopterus Molleri)).

Claes, J. M., & Mallefet, J. (2015). Comparative control of luminescence in sharks: New insights from the slendertail lanternshark (Etmopterus Molleri). Journal of Experimental Marine Biology and Ecology, 467, 87–94. https://doi.org/10.1016/j.jembe.2015.03.008


 * The authors discovered important differences in hormonal control of luminescence in Etmopterus spinax and Etmopterus molleri, both members of the Etmopterus genus. The authors investigated the effects of naturally-occurring neurotransmitters and comparable agonists; a nitric oxide (NO) donor molecule of sodium nitroprusside; hormones melatonin (MT), prolactin (PRL), and ɑ-melanocyte-stimulating hormone (ɑMSH); and receptor antagonists (MT and GABA) on the production of light in the tissues of the photophores collected from E. spinax and E. molleri. In contrast to other members of the Etmopterus genus, such as E.spinax, E. molleri required a higher threshold of PRL input than MT to produce luminescence. The authors hypothesized that E. molleri overall had lower responses to hormones controlling the production of luminescence than E. spinax and required a higher threshold to produce light, which may have interesting ecological implications regarding the implementation of luminescence to each species.

Duchatelet, L., Delroisse, J., & Mallefet, J. (2020). Bioluminescence in Lanternsharks: Insight from hormone receptor localization. General and Comparative Endocrinology, 294. https://doi.org/10.1016/j.ygcen.2020.113488


 * Hormonal control of bioluminescence in Etmopterus molleri is regulated by melatonin (MT), adrenocorticotropic hormone (ACTH), and α-melanocyte-stimulating hormone (αMSH) that act on their respective receptors in the shark’s photophores. In E. molleri, MT and PRL stimulate light production while ACTH and αMSH prevent light production by the photophore. G-protein-coupled receptors, a family of receptors belonging to the melanocortin group, called MCR that may be responsive to both αMSH and ACTH were identified within the photophore The researchers collected samples of tissue from the photophores of E. molleri and were able to localize melatonin receptors (MTNR) and MCR within the tissue of the photophore, as well as demonstrate the effect that their corresponding hormones had on light production.

Duchatelet, L., Delroisse, J., Pinte, N., Sato, K., Ho, H. C., & Mallefet, J. (2019). Adrenocorticotropic hormone and cyclic adenosine monophosphate are involved in the control of shark bioluminescence. Photochemistry and Photobiology, 96(1), 37–45. https://doi.org/10.1111/php.13154


 * The researchers demonstrated that within the Etmopteridae family, adrenocorticotropic hormone (ACTH) that is modulated by cyclic adenosine monophosphate (cAMP) inhibits light emission. ACTH was hypothesized to interact with G-protein-coupled receptors, often identified by the general category MCR, to increase levels of cAMP directly by increasing the amount of adenylate cyclase activity within the cells of the photophore. Specimens of Etmopterus molleri were collected and pharmacological treatments applied to the tissue of the shark’s photophores. The researchers injected ACTH alone to investigate its effect on light emission, followed by a condition where they injected MT followed by ACTH to observe the inhibitory effect of the hormone on light production. In E. molleri, the injection of ACTH following MT effectively decreased light emission as predicted. Additionally, they demonstrated that cAMP concentrations increased following ACTH injection by increasing production of adenylate cyclase. They observed that with MT injection, cAMP concentrations decreased. The researchers hypothesize that the intricate system of moderation of light production that is moderated by cAMP with both inducing/reducing hormones may be especially sensitive in order for the shark to camouflage luminescence when it is beneficial within the aphotic zone of the ocean.

Takahashi, A., & Kawauchi, H. (2006). Evolution of melanocortin systems in fish. General and Comparative Endocrinology, 148(1), 85–94. https://doi.org/10.1016/j.ygcen.2005.09.020


 * This source provides an overview of the evolution of melanocortin system specifically within fish as compared to other invertebrates and vertebrates. Given the importance of the melanocortin system in regulating luminescence in E. molleri, evolutionary differences between the effects of the proteins in different evolutionary lineages suggest that even within the Etmopterus genus each species may have different responsivity to the hormones as evidenced by more recent research. Although the paper does not directly reference E. molleri, I wanted to understand the evolution of melanocortin systems more broadly in fish in order to gain a sense of the diversity of functions that they service in different evolutionary groups. The authors describe how melanocortin molecules in invertebrates were evolutionarily conserved from invertebrates, and extant leech and mussels contain this system as well. Diversification of melanocortin systems in jawed fish may have led to the evolution of a variety of functions by these systems in other fish, which likely is specialized for the control of bioluminescence within the photophores of certain shark families. Although not the focus of my research, melanocortin systems have important evolutionary functions within the vertebrates and the five classes that are most often the subject of research on hormones.