Individual colour patches as multicomponent signals

Biological Reviews of the Cambridge Philosophical Society

Volumn 79, Issue 3, 2004, Pages 583-610

  Grether, Gregory F ^a   Kolluru, Gita R ^a   Nersissian, Karen ^a  

^a UNIVERSITY OF CALIFORNIA   (United States)

Author keywords

Amplifier; Carotenoid; Chromatophore; Colour patch; Handicap; Indicator; Mimicry; Multiple trait; Signal evolution

Indexed keywords

CAROTENOID; MELANIN; PIGMENT; PTERIDINE;

AMPHIBIA; BIOLOGICAL TRAIT; CHROMATOPHORE; COLOR; COLOR PATCH; COMPUTER SIMULATION; ERYTHROPHORE; EVOLUTION; FISH; HABITAT STRUCTURE; IRIDOPHORE; LEUKOPHORE; LIGHT; MATING; MELANOPHORE; NONHUMAN; RECOGNITION; REPTILE; REVIEW; SEXUAL BEHAVIOR; SIGNAL DETECTION; SPECIES DIFFERENTIATION; SPECIES RECOGNITION; THEORY; VERTEBRATE; XANTHOPHORE;

ANIMALS; CHROMATOPHORES; EVOLUTION; MATHEMATICS; MELANOPHORES; MODELS, BIOLOGICAL; PIGMENTATION; SKIN PHYSIOLOGY;

AMPHIBIA; ANIMALIA; MELANOPHORA; PISCES; REPTILIA; VERTEBRATA;

EID: 4344674336     PISSN: 14647931     EISSN: None     Source Type: Journal    
DOI: 10.1017/S1464793103006390     Document Type: Review

References (140)

1. ANDERSSON, M. (1994). Sexual Selection. Princeton University Press, Princeton.
2. ANDERSSON, M. & IWASA, Y. (1996). Sexual Selection. Trends in Ecology & Evolution 11, 53-58.
3. ANDERSSON, S., PRYKE, S. R., ORNBORG, J., LAWES, M. J. & ANDERSSON, M. (2002). Multiple receivers, multiple ornaments, and a trade-off between agonistic and epigamic signaling in a widowbird. The American Naturalist 160, 683-691.
4. ARCHER, S. N. & LYTHGOE, J. N. (1990). The visual pigment basis for cone polymorphism in the guppy, Poecilia reticulata. Vision Research 30, 225-233.
5. BADYAEV, A. V., HILL, G. E., DUNN, P. O. & GLEN, J. C. (2001). Plumage colour as a composite trait: developmental and functional integration of sexual ornamentation. The American Naturalist 158, 221-235.
6. BAGNARA, J. T. (1966). Cytology and cytophysiology of non-melanophore pigment cells. International Review of Cytology 20, 173-205.
7. BAGNARA, J. T. & HADLEY, M. E. (1973). Chromatophores and colour change, the comparative physiology of animal pigmentation. Prentice Hall, Englewood Cliffs.
8. BAGNARA, J. T., TAYLOR, J. D. & HADLEY, M. E. (1968). The dermal chromatophore unit. Journal of Cell Biology 38, 67-79.
9. BENDICH, A. (1989). Carotenoids and the immune system. Journal of Nutrition 119, 112-115.
10. BERGLUND, A., BISAZZA, A. & PILASTRO, A. (1996). Armaments and ornaments: an evolutionary explanation of traits of dual utility. Biological Journal of the Linnean Society 58, 385-399.
11. BLOUNT, J. D., METCALFE, N. B., BIRKHEAD, T. R. & SURAI, P. F. (2003). Carotenoid modulation of immune function and sexual attractiveness in Zebra finches. Science 300, 125-127.
12. BORGIA, G. & PRESGRAVES, D. C. (1998). Coevolution of elaborate male display traits in the spotted bowerbird: an experimental test of the threat reduction hypothesis. Animal Behaviour 56, 1121-1128.
13. BROOKS, R. & COULDRIDGE, V. (1999). Multiple sexual ornaments coevolve with multiple mating preferences. The American Naturalist 154, 37-45.
14. BRUSH, A. H. (1990). Metabolism of carotenoid pigments in birds. The FASEB Journal 4, 2969-2977
15. BURTON, G. W. (1989). Antioxidant actions of carotenoids. The Journal of Nutrition 119, 109-111.
16. CAMPLANI, A., SAINO, N. & MØLLER, A. P. (1999). Carotenoids, sexual signals and immune function in barn swallows from Chernobyl. Proceedings of the Royal Society of London Series B 266, 1111-1116.
17. CHITTKA, L. (1992). The colour hexagon - a chromaticity diagram based on photoreceptor excitations as a generalized representation of colour opponency. Journal of Comparative Physiology a-Sensory Neural and Behavioural Physiology 170, 533-543.
18. CHITTKA, L. (1999). Bees, white flowers, and the colour hexagon - A reassessment? No, not yet: comments on the contribution by Vorobyev et al. Naturwissenschaften 86, 595-597.
19. CONNETT, J. E., KULLER, L. H., KJELSBERG, M. O., POLK, B. F., COLLINS, G., RIDER, A. & HULLEY, S. B. (1989). Relationship between carotenoids and cancer. Cancer 64, 126-134.
20. CRAPON DE CAPRONA, M. D. & RYAN, M. J. (1990). Conspecific mate recognition in swordtails. Xiphophorus nigrensis and X. pygmaeus. (Poeciliidae): olfactory and visual cues. Animal Behaviour 39, 290-296.
21. DARWIN, C. (1871). The Descent of Man, and Selection in Relation to Sex. J. Murray, London.
22. DE IBARRA, N. H., VOROBYEV, M., BRANDT, R. & GIURFA, M. (2000). Detection of bright and dim colours by honeybees. The Journal of Experimental Biology 203, 3289-3298.
23. DENTON, E. J. & LAND, M. F. (1971). Mechanism of reflexion in silvery layers of fish and cephalopods. Proceedings of the Royal Society of London Series A 178, 43-61.
24. ENDLER, J. A. (1980). Natural selection on colour patterns in Poecilia reticulata. Evolution 34, 76-91.
25. ENDLER, J. A. (1991a). Interactions between predators and prey. In Behavioural Ecology, An Evolutionary Approach (eds. J. R. Krebs and N. B. Davies), pp. 169-201. Blackwell, Oxford.
26. ENDLER, J. A. (1991b). Variation in the appearance of guppy colour patterns to guppies and their predators under different visual conditions. Vision Research 31, 587-608.
27. ENDLER, J. A. (1992). Signals, signal conditions, and the direction of evolution. The American Naturalist 139, 1-27.
28. ENDLER, J. A. (1993). The colour of light in forests and its implications. Ecological Monographs 63, 1-27.
29. ENDLER, J.A. & HOUDE, A.E. (1995). Geographic variation in female preferences for male traits in Poecilia reticulata. Evolution 49, 456-468.
30. ENDLER, J. A. & THÉRY, M. (1996). Interacting effects of lek placement, display behaviour, ambient light, and colour patterns in three neotropical forest-dwelling birds. The American Naturalist 148, 421-452.
31. FAIVRE, B., GRÉGOIRE, A., PRÉAULT, M., CÉZILLY, F. & SORGI, G. (2003). Immune activation rapidly mirrored in a seconday sexual trait. Science 300, 103.
32. FISHER, R. A. (1930). The Genetical Theory of Natural Selection. Clarendon Press, Oxford.
33. FISHER, R. A. (1958). The Genetical Theory of Natural Selection. Dover, New York.
34. FITPATRICK, S. (1998a). Birds' tails as signaling devices: markings, shape, length, and feather quality. The American Naturalist 151, 157-173.
35. FITPATRICK, S. (1998b). Colour schemes for birds: structural coloration and signals of quality in feathers. Annales Zoologici Fennici 35, 67-77.
36. FLEISHMAN, L. J., BOWMAN, M., SAUNDERS, D., MILLLER, W. E., RURY, M. J. & LOEW, E. R. (1997). The visual ecology of Puerto Rican anoline lizards: habitat light and spectral sensitivity. Journal of Comparative Physiology a-Sensory Neural and Behavioural Physiology 181, 446-460.
37. FLEISHMAN, L. J. & PERSONS, M. (2001). The influence of stimulus and background colour on signal visibility in the lizard Anolis cristatellus. The Journal of Experimental Biology 204, 1559-1575.
38. FUJII, R. (1993). Cytophysiology of fish chromatophores. International Review of Cytology 143, 191-255.
39. FUJII, R. (2000). The regulation of motile activity in fish chromatophores. Pigment Cell Research 13, 300-319.
40. FUJII, R., HAYASHI, H. & TOYOHARA, J. (1991). Analysis of the reflection of light from motile iridophores of the Dark Sleeper, Odontobutis obscura obscura. Zoological Science 8, 461-470.
41. GOODWING, T. W. (1984). The Biochemistry of the Carotenoids. Chapman & Hall, London.
42. GOODWIN, T. W. (1986). Metabolism, nutrition, and function of carotenoids. Annual Review of Nutrition 6, 274-297.
43. GRAFEN, A. (1990). Sexual selection unhandicapped by the Fisher process. Journal of Theoretical Biology 144, 473-516.
44. GRETHER, G. F. (1996). Intrasexual competition alone favors a sexually dimorphic ornament in the rubyspot damselfly Hetaerina americana. Evolution 50, 1949-1957.
45. GRETHER, G. F. (2000). Carotenoid limitation and mate preference evolution: a test of the indicator hypothesis in guppies (Poecilia reticulata). Evolution 54, 1712-1724.
46. GRETHER, G. F., HUDON, J. & ENDLER, J. A. (2001a). Carotenoid scarcity, synthetic pteridine pigments and the evolution of sexual coloration in guppies (Poecilia reticulata). Proceedings of the Royal Society of London Series B 268, 1245-1253.
47. GRETHER, G. F., HUDON, J. & MILLIE, D. F. (1999). Carotenoid limitation of sexual coloration along an environmental gradient in guppies. Proceedings of the Royal Society of London Series B 266, 1317-1322.
48. GRETHER, G. F., KASAHARA, S., KOLLURU, G. R. & COOPER, E. L. (in press). Sex-specific effects of carotenoid intake on the immunological response to allografts in guppies (Poecilia reticulata). Proceedings of the Royal Society of London Series B.
49. GRETHER, G. F., MILLIE, D. F., BRYANT, M. J., REZNIK, D. N. & MAYEA, W. (2001b). Rain forest canopy cover, resource availability, and life history evolution in guppies. Ecology 82, 1546-1559.
50. HANKINSON, S. J. & MORRIS, M. R. (2002). Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences. Behavioural Ecology and Sociobiology 51, 140-145.
51. HASSON, O. (1989). Amplifiers and the handicap principle in sexual selection: a different emphasis. Proceedings of the Royal Society of London Series B 234, 383-406.
52. HASSON, O. (1990). The role of amplifiers in sexual selection: an intergration of the amplifying and the Fisherian mechanisms. Evolutionary Ecology 4, 277-289.
53. HASSSON, O. (1991). Sexual displays as amplifiers: practical examples with an emphasis on feather decorations. Behavioural Ecology 2, 189-197.
54. HASSON, O. (1997). Towards a general theory of biological signaling. Journal of Theoretical Biology 185, 139-156.
55. HERRING, P. J. (1994). Reflective systems in aquatic animals. Comparative Biochememisty and Physiology 109A, 513-546.
56. HILL, G. E. (1996). Redness as a measure of the production cost of ornamental coloration. Ethology Ecology & Evolution 8, 157-175.
57. HILL, G. E. (1999). Is there an immunological cost to carotenoid-based ornamental coloration? The American Naturalist 154, 589-595.
58. HILL, G. E. (2000). Energetic constraints on expression of carotenoid-based plumage coloration. Journal of Avian Biology 31, 559-566.
59. HOLLAND, B. & RICE, W. R. (1998). Perspective: chase-away sexual selection: antagonistic seduction versus resistance. Evolution 52, 1-7.
60. HOUDE, A. E. & ENDLER, J. A. (1990). Correlated evolution of female mating preferences and male colour patterns in the Guppy Poecilia reticulata. Science 248, 1405-1408.
61. HOUDE, A. E. & TORIO, A. J. (1992). Effect of parasitic infection on male colour pattern and female choice in guppies, Behavioural Ecology 3, 346-351.
62. HUDON, J. (1991). Unusual carotenoid use by the Western tanager (Piranga ludoviciana) and its evolutionary implications. Canadian Journal of Zoology 69, 2311-2320.
63. HUDON, J. (1994). Biotechnological applications of research on animal pigmentation. Biotechnology Advances 12, 49-69.
64. HUDON, J., GRETHER, G. F. & MILLIE, D. F. (in press). Marginal carotenoid differentiation between the sexual and general pigmentation of guppies (Poecilia reticulata) and a possible visual explanation. Physiological and Biochemical Zoology.
65. HURST, D. T. (1980). An Introduction to the Chemistry and Biochemistry of Pyrimidines, Purines, and Pteridines. J. Wiley, New York.
66. HUXLEY, A. F. (1968). A theoretical treatment of the reflexion of light by multilayer structures. The Journal of Experimental Biology 48, 227-245.
67. INOUYE, C. Y., HILL, G. E., STRADI, R. D. & MONTGOMERIE, R. (2001). Carotenoid pigments in male house finch plumage in relation to age, subspecies, and ornamental coloration. The Auk 118, 900-915.
68. IWASA, Y. & POSMIANKOWSKI, A. (1994). The evolution of mate preferences for multiple sexual ornaments. Evolution 48, 853-867.
69. JENNIONS, M. D. & PETRIE, M. (1997). Variation in mate choice and mating preferences: a review of causes and consequences. Biological Reviews of the Cambridge Philosophical Society 72, 283-327.
70. JIROTKUL, M. (1999). Operational sex ratio influences female preference and male-male competition in guppies. Animal Behaviour 58, 287-294.
71. JOHNSTONE, R. A. (1995). Honest advertisement of multiple qualities using multiple signals. Journal of Theoretical Biology 177, 87-94.
72. JOHNSTONE, R. A. (1996). Multiple displays in animal communication: 'backup signals' and 'multiple messages'. Philosophical Transactions of the Royal Society of London B 351, 329-338.
73. KAYSER, H. (1985). Pigments. In Comparative Insect Physiology (eds. G. A. Kerkuts and L. E. Gilbert), pp. 367-415. Academic Press, New York.
74. KELBER, A., VOROBYEV, M. & OSORIO, D. (2003). Animal colour vision - behavioural tests and physiological concepts. Biological Reviews of the Cambridge Philosophical Society 78, 81-118.
75. KODRIC-BROWN, A. (1989). Dietary carotenoids and male mating success in the guppy: an environmental component to female choice. Behavioural Ecology and Sociobiology 25, 393-401.
76. KODRIC-BROWN, A. (1993). Female choice of multiple male criteria in guppies: interacting effects of dominance, coloration and courtship. Behavioural Ecology and Sociobiology 32, 415-420.
77. KODRIC-BROWN, A. & BROWN, J. H. (1984). Truth in advertising: the kinds of traits favored by sexual selection. The American Naturalist 124, 309-323.
78. KOSE, M. & MØLLER, A. P. (1999). Sexual selection, feather breakage and parasites: the importance of white spots in the tail of the barn swallow (Hirundo rustica). Behavioural Ecology and Sociobiology 45, 430-436.
79. KRINSKY, N. I. (1989). Carotenoids and cancer in animal models. Journal of Nutrition 119, 123-126.
80. LAUGHLIN, S. (1981). Neural principles in the peripheral visusal systems of invertebrates. In Invertebrate Visual Centres and Behaviour (ed. H. J. Autrum), pp. 133-280. Springer, New York.
81. LEAL, M. & FLEISHMAN, L. J. (2001). Evidence for habitat partitioning based on adaptation to environmental light in a pair of sympatric lizard species. Proceedings of the Royal Society of London Series B 296, 351-359.
82. LENS, L., VAN DONGEN, S., KARK, S. & KARK, S. & MATTHYSEN, E. (2002). Fluctuating asymmetry as an indicator of fitness: can we bridge the gap between studies? Biological Reviews of the Cambridge Philosophical Society 77, 27-38.
83. Losos, J. B. (1985). An experimental demonstration of the species-recognition role of Anolis dewlap color. Copeia 1985, 905-910.
84. LOZANO, G. A. (1994). Carotenoids, parasites, and sexual selection. Oikos 70, 309-311.
85. LOZANO, G. A. (2001). Carotenoids, immunity, and sexual selection: comparing apples and oranges? The American Naturalist 158, 200-203.
86. MACEDONIA, J. M. (2001). Habitat light, colour variation, and ultraviolet reflectance in the Grand Cayman anole, Anolis conspersus. Biological Journal of the Linnean Society 73, 299-320.
87. MACEDONIA, J. M., BRANDT, Y. & CLARK, D. L. (2002). Sexual dichromatism and differential conspicuousness in two populations of the common collared lizard (Crotaphytus collaris) from Utah and New Mexico, USA. Biological Journal of the Linnean Society 77, 67-85.
88. MACEDONIA, J. M., JAMES, S., WITTLE, L. W. & CLARK, D. L. (2000). Skin pigments and coloration in the Jamaican radiation of Anolis lizards. Journal of Herpetology 34, 99-109.
89. MACEDONIA, J. M. & STAMPS, J. A. (1994). Species recognition in Anolis grahami (Sauria, Iguanidae) - evidence from responses to video playbacks of conspecific and heterospecific displays. Ethology 98, 246-264.
90. MATSUNO, T. (2001). Aquatic animal carotenoids. Fisheries Science 67, 771-783.
91. MCGRAW, K. J., HILL, G. E., STRADI, R. & PARKER, R. S. (2001). The influence of carotenoid acquisition and utilization on the maintenance of species-typical plumage pigmentation in male American goldfinches (Carduelis tristis) and northern cardinals (Cardinalis cardinalis). Physiological and Biochemical Zoology 74, 843-852.
92. MCGRAW, K. J., MACKILLOP, E. A., DALE, J. & HAUBER, M. E. (2002). Different colors reveal different information: how nutritional stress affects the expression of melanin- and structurally based ornamental plumage. The Journal of Experimental Biology 205, 3747-3755.
93. MØLLER, A. P., BIARD, C., BLOUNT, J. D., HOUSTON, D. C., NINNI, P., SAINO, N. & SURAI, P. F. (2000). Carotenoid-dependent signals: indicators of foraging efficiency, immunocompetence or detoxification ability? Avian and Poultry Biology Reviews 11, 137-159.
94. MØLLER, A. P. & POMIANSKI, A. (1993). Why have birds got multiple sexual ornaments? Behavioural Ecology and Sociobiology 32, 167-176.
95. MØLLER, A. P. & THORNHILL, R. (1998). Bilateral symmetry and sexual selection: a meta-analysis. American Naturalist 151, 174-192.
96. MORRISON, R. L. (1995). A transmission electron microscope (TEM) method for determining structural colors reflected by lizard iridophores. Pigment Cell Research 8, 28-36.
97. MORRISON, R. L., RAND, M. S. & FROST-MASON, S. K. (1995). Cellular basis of colour differences in three morphs of the lizard Sceloporus undulatus erythrocheilus. Copeia 1995, 397-408.
98. MORRISON, R. L., SHERBROOKE, W. C. & FROST-MASON, S. K. (1996). Temperature-sensitive, physiologically active iridophores in the lizard Urosaurus ornatus: an ultrastructural analysis of colour change. Copeia 4, 804-812.
99. NICOL, J. A. C. & SOMIYA, H. (1989). The Eyes of Fishes. Oxford University Press, Oxford.
100. NIELSEN, H. I. (1978). Ultrastructural changes in the dermal chromatophore unit of Hyla arborea during colour change. Cell Tissue Research 194, 405-408.
101. NIELSEN, H. J. & DYCK, J. (1978). Adaptation of the tree frog, Hyla cinerea, to colored backgrounds, and the role of the three chromatophore types. Journal of Experimental Zoology 205, 79-94.
102. OBIKA, M. (1988). Ultrastructure and physiological response of leucophores of the medaka Oryzias latipes. Zoological Science 5, 311-321.
103. OLIPHANT, L. W. & HUDON, J. (1993). Pteridines as reflecting pigments and components of reflecting organelles in vertebrates. Pigment Cell Research 6, 205-208.
104. OLSON, V. A, & OWENS, I. P. F. (1998). Costly sexual signals: are carotenoids rare, risky or required? Trends in Ecology and Evolution 13, 510-514.
105. OSHIMA, N. (2001). Direct reception of light by chromatophores of lower vertebrates. Pigment Cell Research 14, 312-319.
106. PAINTER, K. J. (2000). Models for pigment pattern formation in the skin of fishes. Mathematical Models for Biological Pattern Formation 121, 59-82.
107. PALMER, A. R. (1999). Detecting publication bias in meta-analyses: a case study of fluctuating asymmetry and sexual selection. The American Naturalist 154, 220-233
108. PALMER, A. R. (2000). Quasireplication and the contract of error: lessons from sex ratios, heritabilities and fluctuating asymmetry. Annual Review of Ecology and Systematics 31, 441-480.
109. PARSONS, P. A. (1990). Fluctuating asymmetry: an epigenetic measure of stress. Biological Reviews of the Cambridge Philosophical Society 65, 131-145.
110. PERSONS, M. H., FLEISHMAN, L. J., FRYE, M. A. & STIMPHIL, M. E. (1999). Sensory response patterns and the evolution of visual signal design in anoline lizards. Journal of Comparative Physiology A Sensory Neural and Behavioural Physiology 184, 585-607.
111. PFENNIG, K. S. (1998). The evolution of mate choice and the potential for conflict between species and mate-quality recognition. Proceedings of the Royal Society of London Series B 265, 1743-1748.
112. POMIANKOWSKI, A. & IWASA, Y. (1993). Evolution of multiple sexual preferences by Fisher's runaway process of sexual selection. Proceedings of the Royal Society of London Series B 253, 173-181.
113. PRYKE, S. R., ANDERSSON, S. & LAWES, M. J. (2001). Sexual selection of multiple handicaps in the red-collared widowbird: female choice of tail length but not carotenoid display. Evolution 55, 1452-1463.
114. PUTNAM, M. (1992). A review of the nature, function, variability, and supply of pigments in salmonid fish. In Aquaculture and the Environment (eds. N. Depauw and J. Joyce), pp. 245-263. European Aquaculture Society, Gent, Belgium.
115. REMPETERS, G., HENZE, M. & ANDERS, F. (1981). Carotenoids and pteridines in the skin of interspecific hybrids of Xiphophorus. Comparative Biochemistry and Physiology 69B, 91-98.
116. RODIECK, R. W. (1973). The Vertebrate Retina - Priniciples of Structure and Function. Freeman and Company, San Francisco.
117. ROSENTHAL, G. G. & SERVEDIO, M. R. (1999). Chase-away sexual selection: resistance to "resistance". Evolution 53, 296-299.
118. ROWE, C. (1999). Receiver psychology and the evolution of multicomponent signals. Animal Behaviour 58, 921-931.
119. ROWE, C. & GUILFORD, T. (1999). The evolution of multimodal warning displays. Evolutionary Ecology 13, 655-671.
120. RYAN, M. J. (1990). Sexual selection, sensory systems, and sensory exploitation. In Oxford Surveys in Evolutionary Biology (eds. D. Futuyma and J. Antonovics), pp. 157-195. Oxford University Press, Oxford.
121. RYAN, M. J., FOX, J. H., WILCZYNSKI, W. & RAND, A. S. (1990). Sexual selection for sensory exploitation in the frog Physalaemus pustlosus. Nature 343, 66-67.
122. RYAN, M. J. & KEDDY-HECTOR, A. (1992). Directional patterns of female mate choice and the role of sensory biases. The American Naturalist 139, S4-S35.
123. SCHLUTER, D. & PRICE, T. (1993). Honesty, perception and population divergence in sexually selected traits. Proceedings of the Royal Society of London Series B 253, 117-122.
124. STAVENGA, D. G., SMITS, R. P. & HOENDERS, B. J. (1993). Simple exponential functions describing the absorbance bands of visual pigment spectra. Vision Research 33, 1011-1017.
125. STRADI, R., PINI, E. & CELENTANO, G. (2001). Carotenoids in bird plumage: the complement of red pigments in the plumage of wild and captive bullfinch (Pyrrhula pyrrhula). Comparative Biochemistry and Physiology Part B 128, 529-535.
126. SULLIVAN, M. S. (1994). Mate choice as an information gathering process under time constraint: implications for behaviour and signal design. Animal Behaviour 47, 141-151.
127. TAKEUCHI, I. K. (1976). Electron microscopy of two types of reflecting chromatophores (iridophores and leucophores) in guppy, Lebistes reticulatus Peters. Cell Tissue Research 173, 17-27.
128. TAYLOR, J. D. & HADLEY, M. E. (1970). Chromatophores and colour change in the lizard, Anolis carolinensis. Zeitschtift für Zellforschung 104, 282-294.
129. THOMPSON, C. W., HILLGARTH, N., LEU, M. & MCCLURE, H. E. (1997). High parasite load in house finches (Carpodacus mexicanus) is correlated with reduced expression of a sexually selected trait. The American Naturalist 149, 270-294.
130. THOMPSON, C. W. & MOORE, M. C. (1991). Throat colour reliably signals status in male tree lizards, Urosaurus ornatus. Animal Behaviour 42, 745-753.
131. THORNHILL, R., MØLLER, A. P. & GANGESTAD, S. W. (1999). The biological significance of fluctuating asymmetry and sexual selection: a reply to Palmer. American Naturalist 154, 234-241.
132. THORPE, R. S. (2002). Analysis of colour spectra in comparative evolution studies: molecular phylogeny and habitat adaptation in the St. Vincent anole (Anolis trinitatis). Systematic Biology 51, 554-569.
133. VON SCHANTZ, T., BENSCH, S., GRAHN, M., HASSELQUIST, D. & WITTZELL, H. (1999). Good genes, oxidative stress and condition-dependent sexual signals. Proceedings of the Royal Society of London Series B 266, 1-12.
134. VOROBYEV, M. (1999). Evolution of flower colors - a model against experiments: reply to comments by Chittka. Naturwissenschaften 86, 598-600.
135. VOROBYEV, M., HEMPEL DE IBARRA, N., BRANDT, R. & GIURFA, M. (1999). Do "white" and "green" look the same to a bee? Naturwissenchaften 86, 592-594.
136. VOROBYEV, M., OSORIO, D., BENNETT, A. T. D., MARSHALL, N. J. & CUTHILL, I. C. (1998). Tetrachromacy, oil droplets and bird plumage colours. Journal of Comparative Physiology A Sensory Neural and Behavioural Physiology 183, 621-633.
137. WEDEKIND, C., MEYER, P., FRISCHKNECHT, M., NIGGLI, U. A. & PFANDER, H. (1998). Different carotenoids and potential information content of red coloration of male three-spined stickle-back. Journal of Chemical Ecology, 24, 787-801.
138. ZAHAVI, A. (1975). Mate selection - a selection for a handicap. Journal of Theoretical Biology 53, 205-214.
139. ZUK, M., LIGON, J. D. & THORNHILL, R. (1992). Effects of experimental manipulation of male secondary sex characters on female mate preference in jungle fowl. Animal Behaviour 44, 999-1006.
140. ZUK, M., THORNTHILL, R., LIGON, J. D., JOHNSON, K., AUSTAD, S., LIGON, S. H., THORNHILL, N. M. & COSTIN, C. (1990). The role of male ornaments and courtship behaviour in female mate choice of red jungle fowl. The American Naturalist 136, 459-473.