Research People Publications Protocols Links


Carleton KL, Conte MA, Malinsky M, Nandamuri SP, Sandkam B, Meier JI, Mwaiko S, Seehausen O, and Kocher TK. Movement of transposable elements contributes to cichlid diversity. Molecular Ecology. Abstact

Carleton, KL and MR Yourick. Axes of visual adaptation in the ecologically diverse family Cichlidae. Seminars in Cell and Developmental Biology. In press. Abstact Online

Escobar-Camacho D, Carleton KL, Narain DW, Pierotti MER. Visual pigment evolution in Characiformes: the dynamic interplay of teleost whole-genome duplication, surviving opsins and spectral tuning. Molecular Ecology in press. Abstact

Carleton KL, Escobar-Camacho D, Stieb SM, Cortesi F, Marshall NJ. Seeing the rainbow: Mechanisms underlying spectral sensitivity in teleost fishes. Invited review. J Exp Bio 23(Pt 8):jeb193334. Abstact

Sandkam BA, Campello L, O'Brien CS, Nandamuri SP, Gammerdinger WJ, Conte MA, Swaroop A, Carleton KL. Mol Biol Evol 37(7): 2002-14. Tbx2a modulates switching of opsin gene expression. Abstract


Carleton KL, Escobar-Camacho D, and Kocher TD. Visual adaptation could aid sympatric speciation in a deep crater lake. Molecular Ecology 28(23): 5007-5009. Abstract

Stieb SM, deBusserolles F, Carleton KL, Cortesi F, Chung W-S, Dalton B, Hammon LA, Marshall NJ. A detailed investigation of the visual system and visual ecology of the Barrier Reef anemonefish, Amphiprion akindynos. Scientific Reports 9(1) 16459. Abstract DB news

Escobar-Camacho D, Taylor MA, Cheney KL, Green NF, Marshall NJ, and Carleton KL. Color discrimination thresholds in a cichlid fish Metriaclima benetos. J Exp Bio. 222 (17) doi:10.1242/jeb.201160. Abstract

Liebgold E and Carleton KL. The right light: behavior and spectral sensitivity of Eastern tiger salamanders (Ambystoma tigrinum). The Wildlife Society Bulletin. In press.

Yourick MR, Sandkam BA, Gammerdinger WJ, Escobar-Camacho D, Nandamuri SP, Clark FE, Joyce B, Conte MA, Kocher TD, and Carleton KL. Diurnal variation in opsin expression and common housekeeping genes necessitates comprehensive normalization methods for quantitative real-time PCR analyses. Molecular Ecol Resources 19(6) 1447-1460. Abstract

Musilova Z, Cortesi F, Matschiner M, Davies WIL, Patel JS, Stieb SM, de Busserolles F, Malmstrom M, Torresen OK, Brown CJ, Mountford JK, Hanel R, Stenkamp DL, Jakobsen KS, Carleton KL, Jentoft S, Marshall J and Salzburger W. Vision using multiple distinct rod opsins in deep-sea fishes. Science 364: 588-92. Article News & Views UMD CMNS News

Luehrmann M, Carleton KL, Cortesi F, Cheney KL and Marshall NJ. Cardinalfishes (Apogonidae) show visual system adaptations typical of nocturnally and diurnally active fish. Molecular Ecology 28(12) 3025-41. Abstract

Conte MA, Joshi R, Moore EC, Nandamuri SP, Gammerdinger WJ, Roberts RB, Carleton KL, Lien S, Kocher TD. Chromosome-scale assemblies reveal the structural evolution of African cichlid genomes. GigaSciences. 8(4): 1-20. Abstract

Escobar-Camacho D, Pierotti M, Ferenc V, Sharpe DMT, Ramos E, Martins C and Carleton KL. Variable vision in variable environments: the visual system of an invasive cichlid (Cichla monoculus, Agassiz, 1831) in Lake Gatun, Panama. J Exp Biol 222: jeb188300. Abstract UMD news


Nandamuri SP, Conte MA, and Carleton KL. Multiple trans factors and one cis-regulatory deletion contribute to differential expression of opsins in African cichlids. BMC Genomics 19:945. Abstract Link

Luehrmann M, Stieb SM, Carleton KL, Pietzker A, Cheney KL and Marshall NJ. Short-term colour vision plasticity on the reef: Changes in opsin expression under varying light conditions differ between ecologically distinct reef fish species. J Exp Bio in press. Abstract

Sandkam BJ, Dalton BE, Breden F and Carleton KL. Re-viewing guppy color vision: Integrating the molecular and physiological variation in visual tuning of a classic system for sensory drive. Curr Zool in press.


Donohue MW, Carleton KL and Cronin TW. Opsin expression in the central nervous system of the mantis shrimp Neogonodactylus oerstedii. Biol Bull 233(1):58-69..

Nandamuri SP, Yourick MR, and Carleton KL. Adult plasticity in African Cichlids: Rapid changes in opsin expression in response to environmental light differences. Mol Ecol 26 (21): 6036-52. Abstract PDF

Escobar-Camacho D, Marshall NJ, and Carleton KL. Behavioral color vision in a cichlid fish: Metriaclima benetos. J Exp Biology 220: 2887-2899. Abstract CMNS video

Nandamuri SP, Dalton BE and Carleton KL. Determination of the genetic architecture underlying short wavelength sensitivity in Lake Malawi cichlids". J Heredity 108(4): 379-390. Abstract

Stieb SM, Cortesi F , Sueess L, Carleton KL, Salzburger W, and Marshall NJ. Why UV- and red-vision are important in damselfish (Pomacentridae): Structural and expression variation in opsin genes. Molecular Ecology 26:1323-42. Abstract News&Views

Escobar-Camacho D, Ramos, E, Martins C and Carleton KL. The opsin genes of Amazonian cichlids. Molecular Ecology 26: 1343-56. Abstract News&Views Sci American 60s science Science Daily

Dalton BE, de Busserolle F, Marshall NJ and Carleton KL. Retinal specialization through spatially varying cell densities and opsin coexpression in cichlid fish. J Exp Biology 220(Pt 2):266-277. Abstract


Stieb SM, Carleton KL, Cortesi F, Marshall NJ and Salzburger W. Depth dependent plasticity in opsin gene expression varies between damselfish (Pomacentridae) species. Molecular Ecology 25(15): 3645-61. Abstract

Marbach-Ad G, Rietschel CH, Saluja N, Carleton KL and Haag ES. 2016. The Use of Small Group Active Engagement Exercises in an Introductory Biology Course: Benefits, Challenges and Lessons Learned. J Micro and Biol Education 17(3): 360-9. Abstract

Carleton KL, Rietschel CH, and Marbach-Ad G. 2016. Group active engagements using quantitative modeling of physiological concepts in large-enrollment biology classes. J Micro and Biol Education 17(3): 487-9. Abstract

Carleton KL, Dalton BE, Escobar-Camacho D, Nandamuri SP. Proximate and ultimate causes of variable visual sensitivities: insights from cichlid fish radiations. Genesis 54(6): 299-325. Abstract

Phillips GAC, Carleton KL and Marshall NJ. Multiple genetic mechanisms contribute to visual sensitivity variation in the Labridae. Mol Biol Evol 33: 201-15. Abstract


Escobar-Camacho D and Carleton KL. Sensory modalities in cichlid fish behavior. Current Opinion in Behavioral Sciences 6: 115-124. Abstract

Dalton BE, Lu J, Leips J, Cronin TW and Carleton KL. Variable light environments induce plastic spectral tuning by regional opsin coexpression in the African cichlid fish, Metriaclima zebra. Molecular Ecology 24(16): 4193-204. Abstract

Marshall J, Carleton KL, and Cronin T. Colour vision in marine organisms. Curr Op Neurobiol 34C:86-94. Abstract

Cortesi F, Musilova Z, Stieb SM, Hart NS, Siebeck UE, Malmstrom M, Torresen OK, Jentoft S, Cheney KL, Marshall NJ, Carleton KL and Salzburger W. Ancestral duplications and highly dynamic opsin gene evolution in percomorph fishes. PNAS 112: 1493-8. Abstract


Dalton B, Loew ER, Cronin TW and Carleton KL. Spectral tuning by opsin coexpression in retinal regions that view different parts of the visual field. Proceedings of the Royal Society B: Biological Sciences 281: 20141980. Abstract PDF Supplementary methods Supplementary tables

Carleton KL. 2014. Visual photopigment evolution in speciation. In The Evolution of Visual and Non-visual pigments. DM Hunt and M Hankins eds. Springer, New York, pp 241-267.

Brawand D et al (75 authors). The genomic substrate for adaptive radiation in African cichlid fish. Nature 513: 375-81. Abstract

Schulte JE, O'Brien CS, Conte MA, O'Quin KE and Carleton KL. Interspecific variation in Rx1 expression controls opsin expression and causes visual system diversity in African cichlid fishes. Mol Biol Evol 31(9):2297-2308. Abstract PDF Supplementary info Supp Table S2

Redish EF, Bauer C, Carleton KL, Cooke TJ, Cooper M, Crouch CH, Dreyfus BW, Geller B, Giannini J, Svoboda Gouvea J, Klymkowsky MW, Losert W, Moore K, Presson J, Sawtelle V, Thompson KV, & Turpen C. NEXUS/Physics: An interdisciplinary repurposing of physics for biologists. Am J Physics 82: (5) 368. Abstract PDF


Nikaido M, Suzuki H, Toyoda A, Fujiyama A, Hagino-Yamagishi K, Kocher TD, Carleton K and Okada N. 2013. Lineage-specific expansion of vomeronasal type 2 receptor-like (OlfC) genes in cichlids may contribute to diversification of amino acid detection systems. Genome Biol Evol 5(4) 711-22. Abstract


O'Quin KO, Schulte JE, Patel Z, Kahn N, Naseer Z, Wang H, Conte MA and Carleton KL. Evolution of cichlid vision via trans-regulatory divergence. BMC Evol Biol 12:251. Abstract PDF Supplementary info

Hofmann CM, Marshall NJ, Abdilleh K, Patel Z, Siebeck U, and Carleton KL. Opsin evolution in damselfish: Convergence, reversal, and parallel evolution across tuning sites. J Mol Evol 75: 79-91. Abstract PDF Supplementary info

Cahill KB, Quade JH, Carleton KL and Cote RH. Identification of amino acid residues responsible for the selectivity of Tadalafil binding to two closely related phosphodiesterases, PDE5 and PDE6. J Biol Chem 287:41406-16. Abstract

Smith AR, van Staaden MJ and Carleton KL. An evaluation of the role of sensory drive in the evolution of Lake Malawi cichlid fishes. Int J Evol Biol paper 647420. Abstract PDF

Coyle BJ, Hart NS, Carleton KL and Borgia G. Limited variation in visual sensitivity among bowerbird species suggests that there is no link between spectral tuning and variation in display colouration. J Exp Biol 215:1090-1105. Abstract PDF

Smith AR, Ma K, Soares D, and Carleton KL. Relative LWS cone opsin expression determines optomotor thresholds in Malawi cichlid fish. Genes, Brain and Behavior 11: 185-92. Abstract PDF Supplementary info


Carleton KL. Quantification of transcript levels with quantitative RT-PCR. In Molecular methods for evolutionary genetics. Methods in Molecular Biology. V Orgogozo and M Rockman eds. Abstract Scanned pdf

O'Quin KE, Smith D, Naseer Z, Schulte J, Engel SD, Loh Y-H, Streelman JT, Boore JL and Carleton KL. Divergence in cis-regulatory sequences surrounding the opsin gene arrays of African cichlid fishes. BMC Evol Biology 11:120. Abstract PDF

Lesser MP, Carleton KL, Bottger SA, Barry TM and Walker CW. Sea urchin tube feet are photosensory organs that express a rhabdomeric-like opsin and PAX6. Proc Biol Sci 278: 3371-9.

O'Quin KE, Smith AR, Sharma A and Carleton KL. New evidence for the role of heterochrony in the repeated evolution of cichlid opsin expression. Evolution and Development 13:193-203. Abstract PDF

Smith AR, D'Annunzio LD, Smith AE, Sharma A, Hofmann CM, Marshall NJ and Carleton KL. Intraspecific cone opsin expression variation in the cichlids of Lake Malawi. Molecular Ecology 20:299-310. Abstract PDF Supplementary figures Supplementary tables


O'Quin KE, Hofmann CM, Hofmann HA, and Carleton KL. Parallel evolution of opsin gene expresson in African cichlid fishes. Molecular Biology and Evolution 27: 2839-54. Abstract PDF Supplementary info

Smith AR and Carleton KL. Allelic variation in Malawi cichlid fish opsins: a tale of two genera. Journal of Molecular Evolution 70: 593-604. Abstract PDF Supplementary tables

Dalton BE, Cronin TW, Marshall NJ, and Carleton KL. The fish eye view: Are cichlids conspicuous? Journal of Experimental Biology 213: 2243-55. Abstract PDF Supplementary Info

Hofmann CM, O'Quin KE, Smith A and Carleton KL. Plasticity of opsin gene expression in cichlids from Lake Malawi. Molecular Ecology 19: 2064-74. Abstract PDF

Lee B-Y, Howe AE, Conte MA, D'Cotta H, Pepey E, Baroiller J-F, DiPalma F, Carleton KL, and Kocher TD. An EST resource for tilapia based on 17 normalized libraries and assembly of 116,899 sequence tags. BMC Genomics 11:278. PDF

Carleton KL, Hofmann CM, Klisz C, Patel Z, Chircus LM, Simenauer LH, Soodoo N, Albertson RC and Ser JR. Genetic basis of differential opsin gene expression in cichlid fishes. Journal of Evolutionary Biology 23: 840-53. Abstract PDF Supplementary info

Hofmann CM, O'Quin KE, Marshall NJ and Carleton KL. The relationship between lens transmission and opsin gene expression in cichlids from Lake Malawi. Vision Research 50: 357-63. PDF Supplementary info


Hofmann CM*, O'Quin KE*, Marshall NJ, Cronin TW, Seehausen O, and Carleton KL. The eyes have it: Regulatory and structural changes both underlie cichlid visual pigment diversity. PLOS Biology 7(12) e1000266. PDF Synopsis Supplementary Info PLoS blog: Story behind the science

Hofmann CM and Carleton KL. Gene duplication and differential gene expression play an important role in the diversification of visual pigments in fish. Integrative and Comparative Biology 49: 630-43. PDF.

Carleton KL. Cichlid fish visual systems: mechanisms of spectral tuning. Integrative Zoology 4: 75-86. PDF


Seehausen O, Terai Y, Magalhae IS, Carleton KL, Mrosso HDJ, Miyagi R, van der Sluijs I, Schneider MV, Maan ME, Tachida H, Imai H and Okada N. Speciation through sensory drive in cichlid fish. Nature 455: 620-6. Article Nature N&V Supplementary info Other news

Carleton KL, Spady TC, Streelman JT, Kidd MR, Mcfarland WN, and Loew ER. Visual sensitivities tuned by heterochronic shifts in opsin gene expression. BMC Biology 6:22. PDF Supplementary files Mini review


Spady TC, Parry JWL, Robinson PR, Hunt DM, Bowmaker JK and Carleton KL. Evolution of the cichlid visual palette through ontogenetic subfunctionalization of the opsin gene arrays. Mol Biol Evol 23: 1538-1547. PDF

Carleton KL, Spady TC and Kocher TD. Visual communication in East African cichlid fishes : Diversity in a phylogenetic context. In Communication in Fishes. F. Ladich, S.P. Collin, P. Moller and B.G. Kapoor eds. Science Publisher Inc, Enfield, New Hampshire, pp. 481-511. Scanned pdf


Carleton KL, Parry JWL, Bowmaker JK, Hunt DM and Seehausen O. Color vision and speciation in Lake Victoria cichlids of the genus Pundamilia. Molecular Ecology 14: 4341-53. PDF Supplementary table

Parry JWL, Carleton KL, Spady TC, Carboo A, Hunt DM and Bowmaker JK. Mix and match colour vision: tuning spectral sensitivity by differential opsin gene expression in Lake Malawi cichlids. Curr Biol 15: 1734-9. PDF Supplementary Data Commentary Press release

Carleton KL, Spady TC, and Cote RH. Rod and cone opsin families differ in spectral tuning domains but not signal transducing domains as judged by saturated evolutionary trace analysis. J Mol Evol 61: 75-89. PDF Supplementary Data

Katagiri T, Kidd C, Tomasina E, Davis JT, Wishon C, Stern JE, Carleton KL, Howe AE and Kocher TD. A BAC-based physical map of the Nile tilapia genome. BMC Genomics 6:89. PDF

Fuller RC, Carleton KL, Fadool JM, Spady TC and Travis J. Genetic and environmental variation in the visual properties of bluefin killifish, Lucania goodei. J Evol Biol 18: 516-523. PDF

Lee BY, Lee WJ, Streelman JT, Carleton KL, Howe AI, Hulata G, Slettan A, Stern JE, Terai Y and Kocher TD. A second generation genetic linkage map of tilapia (Oreochromis spp.) Genetics 170: 237-244. PDF

Scott MP, Panaitof SC, and Carleton KL. Quantification of vitellogenin-mRNA during maturation and breeding of a burying beetle. J Insect Phys 51: 323-31. PDF

Spady TC, Seehausen O, Loew ER, Jordan RC, Kocher TD and Carleton KL. Adaptive molecular evolution in the opsin genes of rapidly speciating cichlid species. Mol Biol Evol 22: 1412-22. PDF

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