266:
shallow water, this dark tint changed into one of a yellowish green. The colour, examined more carefully, was a French grey, with numerous minute spots of bright yellow: the former of these varied in intensity; the latter entirely disappeared and appeared again by turns. These changes were effected in such a manner that clouds, varying in tint between a hyacinth red and a chestnut-brown, were continually passing over the body. Any part, being subjected to a slight shock of galvanism, became almost black: a similar effect, but in a less degree, was produced by scratching the skin with a needle. These clouds, or blushes as they may be called, are said to be produced by the alternate expansion and contraction of minute vesicles containing variously coloured fluids.
694:
3028:
716:
chromatophores are exposed to light and the skin takes on their hue. Likewise, after melanin aggregation in DCUs, the skin appears green through xanthophore (yellow) filtering of scattered light from the iridophore layer. On the dispersion of melanin, the light is no longer scattered and the skin appears dark. As the other biochromatic chromatophores are also capable of pigment translocation, animals with multiple chromatophore types can generate a spectacular array of skin colours by making good use of the divisional effect.
3570:
3564:
3558:
876:
468:
626:
712:, the dermal melanophores tend to be flat and cover a large surface area. However, in animals with thick dermal layers, such as adult reptiles, dermal melanophores often form three-dimensional units with other chromatophores. These dermal chromatophore units (DCU) consist of an uppermost xanthophore or erythrophore layer, then an iridophore layer, and finally a basket-like melanophore layer with processes covering the iridophores.
276:
527:
38:
1058:
146:
862:
the hue of the immediate environment. It has been demonstrated that the background adaptation process is vision-dependent (it appears the animal needs to be able to see the environment to adapt to it), and that melanin translocation in melanophores is the major factor in colour change. Some animals,
391:
to reach the eye of the observer. Structural colours are produced by various combinations of diffraction, reflection or scattering of light from structures with a scale around a quarter of the wavelength of light. Many such structures interfere with some wavelengths (colours) of light and transmit
1099:
can operate chromatophores in complex, undulating chromatic displays, resulting in a variety of rapidly changing colour schemata. The nerves that operate the chromatophores are thought to be positioned in the brain in a pattern isomorphic to that of the chromatophores they each control. This means
613:
Humans have only one class of pigment cell, the mammalian equivalent of melanophores, to generate skin, hair, and eye colour. For this reason, and because the large number and contrasting colour of the cells usually make them very easy to visualise, melanophores are by far the most widely studied
662:
biochrome of unknown chemical structure in cells named cyanophores. Although they appear unusual in their limited taxonomic range, there may be cyanophores (as well as further unusual chromatophore types) in other fish and amphibians. For example, brightly coloured chromatophores with undefined
265:
These animals also escape detection by a very extraordinary, chameleon-like power of changing their colour. They appear to vary their tints according to the nature of the ground over which they pass: when in deep water, their general shade was brownish purple, but when placed on the land, or in
715:
Both types of melanophore are important in physiological colour change. Flat dermal melanophores often overlay other chromatophores, so when the pigment is dispersed throughout the cell the skin appears dark. When the pigment is aggregated toward the centre of the cell, the pigments in other
867:, have a highly developed background adaptation response capable of generating a number of different colours very rapidly. They have adapted the capability to change colour in response to temperature, mood, stress levels, and social cues, rather than to simply mimic their environment.
580:. When this protein is defective, no melanin can be generated resulting in certain types of albinism. In some amphibian species there are other pigments packaged alongside eumelanin. For example, a novel deep (wine) red-colour pigment was identified in the melanophores of
902:. These cells have the ability to migrate long distances, allowing chromatophores to populate many organs of the body, including the skin, eye, ear, and brain. Fish melanophores and iridophores have been found to contain the smooth muscle regulatory proteins and
747:
control or both and for many species of bony fishes it is known that chromatophores can respond directly to environmental stimuli like visible light, UV-radiation, temperature, pH, chemicals, etc. Neurochemicals that are known to translocate pigment include
974:
Chromatophores are sometimes used in applied research. For example, zebrafish larvae are used to study how chromatophores organise and communicate to accurately generate the regular horizontal striped pattern as seen in adult fish. This is seen as a useful
1008:
of blindness in cold-blooded species, as animals with certain visual defects fail to background adapt to light environments. Human homologues of receptors that mediate pigment translocation in melanophores are thought to be involved in processes such as
2738:
cuttlefish have instead relied on invisibility, a talent that may have applications for human technology. Norman said the military has shown interest in cuttlefish camouflage with a view to one day incorporating similar mechanisms in soldiers'
152:
151:
148:
147:
439:
containing pteridine and carotenoids are sometimes found in the same cell, in which case the overall colour depends on the ratio of red and yellow pigments. Therefore, the distinction between these chromatophore types is not always clear.
153:
834:
1084:, and sheath cells. Inside the chromatophore cell, pigment granules are enclosed in an elastic sac, called the cytoelastic sacculus. To change colour the animal distorts the sacculus form or size by muscular contraction, changing its
1080:) have complex multicellular organs that they use to change colour rapidly, producing a wide variety of bright colours and patterns. Each chromatophore unit is composed of a single chromatophore cell and numerous muscle, nerve,
479:. When illuminated they generate iridescent colours because of the constructive interference of light. Fish iridophores are typically stacked guanine plates separated by layers of cytoplasm to form microscopic, one-dimensional,
150:
1092:. This differs from the mechanism used in fish, amphibians, and reptiles in that the shape of the sacculus is changed, rather than translocating pigment vesicles within the cell. However, a similar effect is achieved.
142:(blue). While most chromatophores contain pigments that absorb specific wavelengths of light, the color of leucophores and iridophores is produced by their respective scattering and optical interference properties.
918:
and the neural tube. The exception to this is the melanophores of the retinal pigmented epithelium of the eye. These are not derived from the neural crest. Instead, an outpouching of the neural tube generates the
2609:
Jayawickreme, CK; Sauls, H; Bolio, N; Ruan, J; Moyer, M; Burkhart, W; Marron, B; Rimele, T; Shaffer, J (1999). "Use of a cell-based, lawn format assay to rapidly screen a 442,368 bead-based peptide library".
415:(erythrocytes), which are generated in bone marrow throughout the life of an organism, rather than being formed during embryological development. Therefore, erythrocytes are not classified as chromatophores.
514:
shine. As with xanthophores and erythrophores, in fish the distinction between iridophores and leucophores is not always obvious, but, in general, iridophores are considered to generate iridescent or
483:. Both the orientation and the optical thickness of the chemochrome determines the nature of the colour observed. By using biochromes as coloured filters, iridophores create an optical effect known as
610:. While it is likely that other lesser-studied species have complex melanophore pigments, it is nevertheless true that the majority of melanophores studied to date do contain eumelanin exclusively.
360:
It was only in the 1960s that chromatophores were well enough understood to enable them to be classified based on their appearance. This classification system persists to this day, even though the
2558:
Lamason, RL; Mohideen, MA; Mest, JR; Wong, AC; Norton, HL; Aros, MC; Jurynec, MJ; Mao, X; et al. (2005). "SLC24A5, a putative cation exchanger, affects pigmentation in zebrafish and humans".
1685:
Goda M, Ohata M, Ikoma H, Fujiyoshi Y, Sugimoto M, Fujii R (2011). "Integumental reddish-violet coloration owing to novel dichromatic chromatophores in the teleost fish, Pseudochromis diadema".
193:
and may be initiated by changes in mood, temperature, stress or visible changes in the local environment. Chromatophores are studied by scientists to understand human disease and as a tool in
459:. The absence of carotene in the frogs' diet meant that the red/orange carotenoid colour 'filter' was not present in their erythrophores. This made the frogs appear blue instead of green.
590:, use melanocytes in response to certain signals and hormonal changes, and is capable of becoming colors ranging from bright blue, brown, and black. This was subsequently identified as
2122:
Valverde, P; Healy, E; Jackson, I; Rees, JL; Thody, AJ (1995). "Variants of the melanocyte-stimulating hormone receptor gene are associated with red hair and fair skin in humans".
1247:
Journal Of
Researches Into The Natural History And Geology Of The Countries Visited During The Voyage Round The World Of H.M.S. 'Beagle' Under The Command Of Captain Fitz Roy, R.N
2087:
Logan, DW; Bryson-Richardson, RJ; Pagán, KE; Taylor, MS; Currie, PD; Jackson, IJ (2003). "The structure and evolution of the melanocortin and MCH receptors in fish and mammals".
161:
Some species can rapidly change colour through mechanisms that translocate pigment and reorient reflective plates within chromatophores. This process, often used as a type of
2645:
Andersson, TP; Filippini, D; Suska, A; Johansson, TL; Svensson, SP; Lundström, I (2005). "Frog melanophores cultured on fluorescent microbeads: biomimic-based biosensing".
1104:. This may explain why, as the neurons are activated in iterative signal cascade, one may observe waves of colour changing. Like chameleons, cephalopods use physiological
2722:
1777:
Palazzo, RE; Lynch, TJ; Lo, SJ; Taylor, JD; Tchen, TT (1989). "Rearrangements of pterinosomes and cytoskeleton accompanying pigment dispersion in goldfish xanthophores".
1223:
Sangiovanni, G (1819). "Descrizione di un particolare sistema di organi cromoforo espansivo-dermoideo e dei fenomeni che esso produce, scoperto nei molluschi cefaloso".
510:(often guanine) to reflect light. Unlike iridophores, leucophores have more organized crystals that reduce diffraction. Given a source of white light, they produce a
1919:
Aspengren, S; Sköld, HN; Quiroga, G; Mårtensson, L; Wallin, M (2003). "Noradrenaline- and melatonin-mediated regulation of pigment aggregation in fish melanophores".
2165:
Richardson, J; Lundegaard, PR; Reynolds, NL; Dorin, JR; Porteous, DJ; Jackson, IJ; Patton, EE (2008). "mc1r
Pathway regulation of zebrafish melanosome dispersion".
942:
each of the cell classes found in the adult fish—melanophores, xanthophores and iridophores—are already present. Studies using mutant fish have demonstrated that
962:
are important in controlling chromatophore differentiation. If these proteins are defective, chromatophores may be regionally or entirely absent, resulting in a
149:
3328:
2952:
854:
Most fish, reptiles and amphibians undergo a limited physiological colour change in response to a change in environment. This type of camouflage, known as
560:
because of its light absorbing qualities. It is packaged in vesicles called melanosomes and distributed throughout the cell. Eumelanin is generated from
776:
that, in turn, transduce the signal into the cell. Melanocortins result in the dispersion of pigment, while melatonin and MCH results in aggregation.
701:
Many species are able to translocate the pigment inside their chromatophores, resulting in an apparent change in body colour. This process, known as
226:
The octopus ... seeks its prey by so changing its colour as to render it like the colour of the stones adjacent to it; it does so also when alarmed.
1992:
Logan, DW; Bryson-Richardson, RJ; Taylor, MS; Currie, P; Jackson, IJ (2003). "Sequence characterization of teleost fish melanocortin receptors".
1305:
Animal
Biochromes and Structural Colors: Physical, Chemical, Distributional & Physiological Features of Colored Bodies in the Animal World.
735:
The control and mechanics of rapid pigment translocation has been well studied in a number of different species, in particular amphibians and
3136:
1513:
Ito, S; Wakamatsu, K. (2003). "Quantitative analysis of eumelanin and pheomelanin in humans, mice, and other animals: a comparative review".
1280:
447:, but xanthophores appear to have supplemental biochemical pathways enabling them to accumulate yellow pigment. In contrast, carotenoids are
1436:
Morrison, RL. (1995). "A transmission electron microscopic (TEM) method for determining structural colors reflected by lizard iridophores".
3027:
2920:
938:) develop into their daughter subtypes is an area of ongoing research. It is known in zebrafish embryos, for example, that by 3 days after
3613:
768:(MCH), that are produced mainly in the pituitary, pineal gland, and hypothalamus, respectively. These hormones may also be generated in a
475:
Iridophores, sometimes also called guanophores, are chromatophores that reflect light using plates of crystalline chemochromes made from
708:, is most widely studied in melanophores, since melanin is the darkest and most visible pigment. In most species with a relatively thin
3255:
1204:
980:
2758:
1389:
1312:
720:
3569:
3563:
3321:
3202:
693:
3557:
2945:
2733:
802:
31:
353:) meaning "cell") was adopted for the cells responsible for colour found in birds and mammals. Only one such cell type, the
2863:
Demski, LS (1992). "Chromatophore systems in teleosts and cephalopods: a levels oriented analysis of convergent systems".
765:
3598:
1108:. They are also among the most skilled at camouflage, having the ability to match both the colour distribution and the
906:. Leaving the neural crest in waves, chromatophores take either a dorsolateral route through the dermis, entering the
3930:
3314:
3064:
2727:
2682:"The melanophore aggregating response of isolated fish scales: a very rapid and sensitive diagnosis of whooping cough"
1904:
Meyer-Rochow, VB (2001). "Fish chromatophores as sensors of environmental stimuli". In Kapoor BG & Hara TJ (ed.).
480:
1548:
Bagnara, J.T.; Taylor, JD; Prota, G (1973). "Color changes, unusual melanosomes, and a new pigment from leaf frogs".
3260:
2938:
1150:
Scott M. Boback & Lynn M. Siefferman (2010). "Variation in Color and Color Change in Island and
Mainland Boas (
773:
772:
fashion by cells in the skin. At the surface of the melanophore, the hormones have been shown to activate specific
697:
Fish and frog melanophores are cells that can change colour by dispersing or aggregating pigment-containing bodies.
3800:
3795:
3131:
2680:
Karlsson, JO; Andersson, RG; Askelöf, P; Elwing, H; Granström, M; Grundström, N; Lundström, I; Ohman, L (1991).
2310:"Protein kinase A, which regulates intracellular transport, forms complexes with molecular motors on organelles"
879:
809:
of pigment translocation. Through a mechanism not yet fully understood, cAMP influences other proteins such as
257:
2425:
Meyer-Rochow, VB; Royuela, M (2002). "Calponin, caldesmon, and chromatophores: the smooth muscle connection".
287:. Structural green and blue colours are generated by overlaying chromatophore types to reflect filtered light.
534:
zebrafish larva that fails to synthesise melanin in its melanophores, at the top a non-mutant, wildtype larva
1381:
785:
1634:
Schwalm, PA; Starrett, PH; McDiarmid, RW (1977). "Infrared reflectance in leaf-sitting neotropical frogs".
756:
on the surface on melanophores. The primary hormones involved in regulating translocation appear to be the
451:
and transported to erythrophores. This was first demonstrated by rearing normally green frogs on a diet of
3935:
1855:
Deacon, SW; Serpinskaya, AS; Vaughan, PS; Lopez
Fanarraga, M; Vernos, I; Vaughan, KT; Gelfand, VI (2003).
920:
654:
444:
2361:
3902:
3805:
3265:
3224:
3219:
3197:
1211:
891:
681:
649:
631:
372:
107:
779:
Numerous melanocortin, MCH and melatonin receptors have been identified in fish and frogs, including a
230:
177:, have complex chromatophore organs controlled by muscles to achieve this, whereas vertebrates such as
3426:
3810:
3416:
3245:
3177:
3126:
2827:
2567:
2321:
2272:
2213:
2001:
1643:
1557:
1005:
943:
1957:
Logan, D. W.; Burn, SF; Jackson, IJ (2006). "Regulation of pigmentation in zebrafish melanophores".
411:, for example, is a biochrome responsible for the red appearance of blood. It is found primarily in
399:
Whereas all chromatophores contain pigments or reflecting structures (except when there has been a
3874:
3639:
3529:
3514:
3493:
3478:
2925:
1026:
488:
3453:
3192:
2901:
2796:
2591:
2540:
2450:
2147:
2025:
1837:
1710:
1667:
1581:
1193:
1171:
780:
218:
52:
1401:
Taylor, JD. (1969). "The effects of intermedin on the ultrastructure of amphibian iridophores".
983:
field. Chromatophore biology has also been used to model human condition or disease, including
875:
467:
3743:
3678:
3295:
3290:
3250:
3209:
3157:
2880:
2845:
2788:
2703:
2662:
2627:
2583:
2532:
2491:
2442:
2386:
2339:
2290:
2241:
2182:
2139:
2104:
2066:
2017:
1974:
1936:
1886:
1829:
1794:
1759:
1702:
1659:
1616:
1573:
1530:
1495:
1453:
1418:
1385:
1359:
1308:
1286:
1276:
1121:
1109:
1105:
1042:
1041:
applications of chromatophore-mediated colour changes have been proposed, mainly as a type of
841:
625:
614:
chromatophore. However, there are differences between the biology of melanophores and that of
595:
436:
368:
88:
1376:
Bagnara JT. Comparative
Anatomy and Physiology of Pigment Cells in Nonmammalian Tissues. In:
502:
A related type of chromatophore, the leucophore, is found in some fish, in particular in the
110:. Mature chromatophores are grouped into subclasses based on their colour under white light:
3898:
3869:
3779:
3668:
3658:
3583:
3578:
3488:
3463:
3275:
3108:
3000:
2980:
2872:
2835:
2780:
2693:
2654:
2619:
2575:
2522:
2481:
2434:
2376:
2329:
2280:
2231:
2221:
2174:
2131:
2096:
2056:
2009:
1966:
1928:
1876:
1868:
1821:
1786:
1749:
1741:
1694:
1651:
1608:
1565:
1522:
1487:
1445:
1410:
1349:
1341:
1268:
1163:
1101:
1089:
1022:
810:
806:
744:
664:
607:
456:
384:
364:
of the pigments may be more useful to a scientific understanding of how the cells function.
280:
190:
2816:"Behavioral analysis of cuttlefish traveling waves and its implications for neural control"
3890:
3764:
3736:
3726:
3721:
3709:
3694:
3634:
3539:
3483:
3059:
2995:
2985:
1208:
1034:
858:, most commonly appears as a slight darkening or lightening of skin tone to approximately
814:
581:
569:
564:
in a series of catalysed chemical reactions. It is a complex chemical containing units of
515:
503:
432:
1599:
Bagnara, J.T. (2003). "Enigmas of
Pterorhodin, a Red Melanosomal Pigment of Tree Frogs".
2831:
2571:
2325:
2308:
Kashina, AS; Semenova, IV; Ivanov, PA; Potekhina, ES; Zaliapin, I; Rodionov, VI (2004).
2276:
2217:
2005:
1647:
1561:
886:
showing the dorsolateral (red) and ventromedial (blue) routes of chromatoblast migration
3925:
3831:
3653:
3644:
3618:
3608:
3603:
3468:
3458:
3182:
3098:
3086:
2990:
2698:
2681:
2261:"Functional coordination of microtubule-based and actin-based motility in melanophores"
2013:
1881:
1856:
1754:
1729:
1449:
1354:
1325:
1242:
1018:
958:
636:
586:
484:
412:
304:
248:
194:
182:
2623:
2362:"Behavioral genetic approaches to visual system development and function in zebrafish"
2285:
2260:
2236:
2201:
2100:
1491:
1326:"Studies on fine structure and cytochemical properties of erythrophores in swordtail,
1272:
299:) as the name for pigment-bearing cells derived from the neural crest of cold-blooded
3919:
3406:
3346:
3091:
3069:
3044:
3005:
2967:
2527:
2510:
2202:"Intracellular actin-based transport: How far you go depends on how often you switch"
2061:
2044:
1970:
1932:
1698:
1612:
1526:
1414:
939:
822:
749:
703:
2915:
2800:
2751:
2544:
2454:
2029:
1671:
1585:
1175:
275:
3815:
3714:
3473:
3373:
3103:
3049:
2771:
Cloney, RA; Florey, E (1968). "Ultrastructure of cephalopod chromatophore organs".
2595:
2151:
1841:
1825:
1714:
976:
911:
895:
757:
753:
619:
361:
234:
103:
1569:
1025:
for rapidly identifying potential bioactive compounds using melanophores from the
672:
55:-containing cells, or groups of cells, found in a wide range of animals including
618:. In addition to eumelanin, melanocytes can generate a yellow/red pigment called
30:
This article is about a type of cell or multicellular organ. For other uses, see
3769:
3524:
3401:
3280:
3162:
3081:
3010:
2975:
1126:
1085:
1014:
998:
931:
899:
818:
794:
790:
724:
591:
526:
393:
127:
2658:
2206:
Proceedings of the
National Academy of Sciences of the United States of America
598:
that accumulates around eumelanin core, and it is also present in a variety of
3906:
3861:
3673:
3663:
3509:
3411:
3368:
3337:
3270:
3167:
3015:
2840:
2815:
2404:
2334:
2309:
1812:
Porras, MG; De Loof, A; Breuer, M; Aréchiga, H (2003). "Procambarus clarkii".
1478:
Fujii, R. (2000). "The regulation of motile activity in fish chromatophores".
1096:
1081:
1077:
1062:
1046:
849:
728:
668:
615:
577:
539:
448:
388:
376:
354:
300:
252:
213:
162:
123:
84:
72:
68:
17:
3894:
3836:
3534:
3383:
3378:
3285:
3118:
3074:
3054:
2579:
2226:
1201:
1131:
1069:
1030:
963:
952:
903:
837:
769:
761:
603:
599:
545:
380:
205:
178:
157:
7x speed timelapse video of fish melanophores responding to 200μM adrenaline
99:
95:
56:
37:
27:
Cells with a primary function of coloration found in a wide range of animals
2849:
2666:
2631:
2587:
2536:
2495:
2486:
2469:
2446:
2390:
2343:
2245:
2186:
2108:
2070:
2021:
1978:
1940:
1890:
1833:
1706:
1655:
1620:
1534:
1499:
2884:
2792:
2707:
2294:
2178:
2143:
1872:
1798:
1763:
1577:
1457:
1422:
1363:
1290:
987:
and albinism. Recently, the gene responsible for the melanophore-specific
3759:
3731:
3152:
1790:
1663:
1345:
1038:
1010:
984:
907:
561:
452:
404:
400:
1745:
833:
719:
3649:
3421:
3214:
3187:
2784:
2438:
2135:
1057:
1029:. Other scientists have developed techniques for using melanophores as
993:
915:
740:
736:
573:
549:
476:
209:
186:
174:
64:
2876:
2381:
3774:
3306:
2200:
Snider, J; Lin, F; Zahedi, N; Rodionov, V; Yu, CC; Gross, SP (2004).
1167:
997:, was shown to have a human equivalent that strongly correlates with
924:
801:
is required in zebrafish for dispersion of melanin. Inside the cell,
709:
565:
531:
507:
424:
76:
2930:
1267:. International Review of Cytology. Vol. 20. pp. 173–205.
894:, chromatophores are one of a number of cell types generated in the
396:
by creating different colours when seen from different directions.
3519:
3240:
2470:"Genetic analysis of melanophore development in zebrafish embryos"
1330:, with special reference to their pigment granules (pterinosomes)"
1073:
1056:
883:
874:
864:
859:
832:
718:
692:
648:
Nearly all the vibrant blues in animals and plants are created by
624:
557:
553:
525:
511:
496:
466:
375:
or "schemochromes". The biochromes include true pigments, such as
274:
144:
42:
36:
2045:"Melatonin, melatonin receptors and melanophores: a moving story"
1100:
the pattern of colour change functionally matches the pattern of
659:
492:
408:
80:
60:
3310:
2934:
1033:, and for rapid disease detection (based on the discovery that
94:
Chromatophores are largely responsible for generating skin and
3036:
428:
739:
fish. It has been demonstrated that the process can be under
427:
pteridine pigments are named xanthophores; those with mainly
392:
others, simply because of their scale, so they often produce
1857:"Dynactin is required for bidirectional organelle transport"
1265:
Cytology and cytophysiology of non-melanophore pigment cells
1065:, using background adaptation to mimic the local environment
1037:
blocks pigment aggregation in fish melanophores). Potential
1245:(1860). "Chapter 1. Habits of a Sea-slug and Cuttle-fish".
1192:. IX, 622a: 2-10. About 400 BC. Cited in Luciana Borrelli,
367:
Colour-producing molecules fall into two distinct classes:
844:
on exposure to dark (top) and light environments (bottom).
2773:
2723:"Cuttlefish Change Color, Shape-Shift to Elude Predators"
2259:
Rodionov, VI; Hope, AJ; Svitkina, TM; Borisy, GG (1998).
1908:. Science Publishers Enfield (NH), USA. pp. 317–334.
898:, a paired strip of cells arising at the margins of the
407:), not all pigment-containing cells are chromatophores.
1021:. Therefore, pharmaceutical companies have developed a
51:
are cells that produce color, of which many types are
1378:
The
Pigmentary System: Physiology and Pathophysiology
1112:
of their local environment with remarkable accuracy.
518:, whereas leucophores produce reflective white hues.
2511:"Genetics and evolution of pigment patterns in fish"
3849:
3824:
3788:
3752:
3702:
3691:
3627:
3591:
3577:
3502:
3446:
3439:
3394:
3361:
3354:
3345:
3233:
3145:
3117:
3035:
2966:
2926:
Tree of Life Web Project: Cephalopod Chromatophores
2814:Laan, A; Guthnick, T; Kuba, MJ; Laurent, G (2014).
2043:Sugden, D; Davidson, K; Hough, KA; Teh, MT (2004).
443:Most chromatophores can generate pteridines from
387:that makes up white light while permitting other
383:. These pigments selectively absorb parts of the
1307:University of California Press, Berkeley, 1976.
817:carrying pigment containing vesicles along both
652:rather than by pigments. However, some types of
303:and cephalopods. The word itself comes from the
2921:Video footage of squid chromatophore patterning
2902:"Nature's Palette - how animals produce colour"
435:carotenoids are termed erythrophores. However,
263:
251:described the colour-changing abilities of the
224:
2916:Video footage of octopus background adaptation
576:rings. The key enzyme in melanin synthesis is
506:. Like iridophores, they utilize crystalline
3322:
2946:
1198:A catalogue of body patterning in Cephalopoda
639:stripe with an unusual type of chromatophore.
584:. Some species of anole lizards, such as the
423:Chromatophores that contain large amounts of
239:
8:
1952:
1950:
1728:Bagnara, JT; Taylor, JD; Hadley, ME (1968).
789:, a melanocortin receptor known to regulate
83:, in contrast, have a class of cells called
979:system for understanding patterning in the
348:
341:
335:) meaning "bearing". In contrast, the word
330:
323:
316:
309:
3699:
3588:
3443:
3358:
3351:
3329:
3315:
3307:
2953:
2939:
2931:
1258:
1256:
2839:
2732:. Wellington, New Zealand. Archived from
2697:
2526:
2485:
2380:
2355:
2353:
2333:
2284:
2235:
2225:
2060:
1880:
1753:
1353:
805:(cAMP) has been shown to be an important
797:in humans. It has been demonstrated that
2468:Kelsh, RN; Schmid, B; Eisen, JS (2000).
357:, has been identified in these animals.
2082:
2080:
1473:
1471:
1469:
1467:
1142:
245:in an Italian science journal in 1819.
934:chromatophore precursor cells (called
914:, or a ventromedial route between the
731:photography during pigment aggregation
3137:Evolution of color vision in primates
2752:Nanotubes for noisy signal processing
1017:, making them attractive targets for
295:was adopted (following Sangiovanni's
7:
3609:Flotation devices ("secondary fins")
1106:colour change for social interaction
3901:) → Juvenile → Subadult → Adult •
2699:10.1111/j.1574-6968.1991.tb04860.x
2014:10.1111/j.1749-6632.2003.tb03196.x
1450:10.1111/j.1600-0749.1995.tb00771.x
1200:. Firenze University Press, 2006.
1072:cephalopods (including octopuses,
1004:Chromatophores are also used as a
981:evolutionary developmental biology
25:
2759:University of Southern California
2721:Hansford, Dave (August 6, 2008).
1492:10.1034/j.1600-0749.2000.130502.x
1249:. John Murray, London. p. 7.
102:animals and are generated in the
3645:Hepatopancreas (digestive gland)
3568:
3562:
3556:
3026:
2528:10.1111/j.1600-0749.2004.00174.x
2062:10.1111/j.1600-0749.2004.00185.x
1971:10.1111/j.1600-0749.2006.00307.x
1933:10.1034/j.1600-0749.2003.00003.x
1699:10.1111/j.1755-148X.2011.00861.x
1613:10.1034/j.1600-0749.2003.00075.x
1527:10.1034/j.1600-0749.2003.00072.x
41:Chromatophores in the skin of a
3614:Funnel–mantle locking apparatus
1906:Sensory Biology of Jawed Fishes
1730:"The dermal chromatophore unit"
1049:make objects nearly invisible.
271:Classification of chromatophore
3801:Nuchal folds (occipital folds)
3796:Nuchal crest (occipital crest)
1826:10.1016/j.peptides.2003.08.016
803:cyclic adenosine monophosphate
629:The purple-striped dottyback,
419:Xanthophores and erythrophores
32:Chromatophore (disambiguation)
1:
2865:Brain, Behavior and Evolution
2624:10.1016/S1056-8719(00)00083-6
2286:10.1016/S0960-9822(98)70064-8
2101:10.1016/S0888-7543(02)00037-X
1570:10.1126/science.182.4116.1034
1273:10.1016/S0074-7696(08)60801-3
766:melanin-concentrating hormone
208:mentioned the ability of the
181:generate a similar effect by
3907:Protoconch (embryonic shell)
1415:10.1016/0016-6480(69)90157-9
923:, which, in turn, forms the
471:Leucophore layer composition
3065:Simple eye in invertebrates
2612:J Pharmacol Toxicol Methods
1861:The Journal of Cell Biology
910:through small holes in the
882:of a developing vertebrate
774:G-protein-coupled receptors
663:pigments are found in both
463:Iridophores and leucophores
342:
324:
310:
173:. Cephalopods, such as the
167:physiological colour change
3952:
3261:Infrared sensing in snakes
2659:10.1016/j.bios.2004.08.043
847:
537:
233:was the first to describe
212:to change colour for both
29:
3885:
3554:
3132:Evolution of color vision
3024:
2841:10.1016/j.cub.2014.06.027
2335:10.1016/j.cub.2004.10.003
1687:Pigment Cell Melanoma Res
1053:Cephalopod chromatophores
658:do possess vesicles of a
349:
331:
317:
237:pigment-bearing cells as
321:) meaning "colour," and
258:The Voyage of the Beagle
2580:10.1126/science.1116238
2227:10.1073/pnas.0403092101
1779:Cell Motil Cytoskeleton
1382:Oxford University Press
1225:G. Enciclopedico Napoli
863:such as chameleons and
840:chromatophores mediate
679:have been described in
2487:10.1006/dbio.2000.9840
2360:Neuhauss, SC. (2003).
1656:10.1126/science.860137
1156:Journal of Herpetology
1066:
970:Practical applications
887:
845:
732:
727:melanophore imaged by
698:
675:chromatophores, named
655:Synchiropus splendidus
640:
568:and dihydroxyindole-2-
535:
472:
445:guanosine triphosphate
385:visible light spectrum
288:
268:
240:
228:
216:and signalling in his
185:. Such signals can be
158:
45:
3888:Developmental stages:
3780:Spadix and antispadix
3664:Nephridia ("kidneys")
3266:Monocular deprivation
3225:Underwater camouflage
3220:Structural coloration
3198:Disruptive coloration
2515:Pigment Cell Research
2179:10.1089/zeb.2008.0541
1959:Pigment Cell Research
1873:10.1083/jcb.200210066
1601:Pigment Cell Research
1324:Matsumoto, J (1965).
1060:
944:transcription factors
892:embryonic development
878:
856:background adaptation
842:background adaptation
836:
829:Background adaptation
722:
696:
689:Pigment translocation
682:Pseudochromis diadema
650:structural coloration
632:Pseudochromis diadema
628:
544:Melanophores contain
529:
470:
285:Chamaeleo calyptratus
278:
222:(ca 4th century BC):
156:
108:embryonic development
40:
3246:Blindness in animals
3178:Counter-illumination
3127:Evolution of the eye
2736:on August 10, 2008.
1791:10.1002/cm.970130103
1346:10.1083/jcb.27.3.493
1263:Bagnara, JT (1966).
1196:, Graziano Fiorito.
1045:, which could as in
582:phyllomedusine frogs
3875:Squid giant synapse
2832:2014CBio...24.1737L
2728:National Geographic
2686:FEMS Microbiol Lett
2647:Biosens Bioelectron
2572:2005Sci...310.1782L
2405:"Chameleon Powders"
2326:2004CBio...14.1877K
2277:1998CBio....8..165R
2218:2004PNAS..10113204S
2006:2003NYASA.994..319L
1746:10.1083/jcb.38.1.67
1648:1977Sci...196.1225S
1562:1973Sci...182.1034B
1403:Gen Comp Endocrinol
1328:Xiphophorus helleri
1102:neuronal activation
1095:Octopuses and most
1088:, reflectivity, or
1027:African clawed frog
677:erythro-iridophores
491:, producing bright-
489:Rayleigh scattering
138:(black/brown), and
3931:Cephalopod zootomy
3806:Occipital membrane
3679:Pericardial glands
3374:Argonautid eggcase
3193:Deimatic behaviour
2785:10.1007/BF00347297
2509:Kelsh, RN (2004).
2439:10.1002/jemt.10169
2427:Microsc. Res. Tech
2136:10.1038/ng1195-328
1207:2018-02-06 at the
1194:Francesca Gherardi
1190:Historia Animalium
1067:
991:zebrafish strain,
890:During vertebrate
888:
846:
733:
699:
641:
536:
473:
373:structural colours
289:
231:Giosuè Sangiovanni
219:Historia animalium
159:
46:
3913:
3912:
3845:
3844:
3744:Suckers and hooks
3687:
3686:
3669:Nidamental glands
3599:Dermal structures
3552:
3551:
3548:
3547:
3435:
3434:
3304:
3303:
3296:Visual perception
3291:Underwater vision
3256:Feature detection
3251:Eyespot apparatus
3210:Eyespot (mimicry)
3158:Animal coloration
2961:Vision in animals
2877:10.1159/000113909
2826:(15): 1737–1742.
2382:10.1002/neu.10165
1282:978-0-12-364320-9
1122:Animal coloration
1043:active camouflage
665:poison dart frogs
191:neurotransmitters
154:
16:(Redirected from
3943:
3899:Doratopsis stage
3870:Squid giant axon
3789:Occipital region
3700:
3640:Ctenidia (gills)
3635:Branchial hearts
3628:Internal anatomy
3592:External anatomy
3589:
3572:
3566:
3560:
3444:
3359:
3352:
3331:
3324:
3317:
3308:
3276:Palpebral (bone)
3109:Schizochroal eye
3030:
2955:
2948:
2941:
2932:
2912:
2908:
2906:
2889:
2888:
2860:
2854:
2853:
2843:
2811:
2805:
2804:
2768:
2762:
2748:
2742:
2741:
2718:
2712:
2711:
2701:
2677:
2671:
2670:
2642:
2636:
2635:
2606:
2600:
2599:
2566:(5755): 1782–6.
2555:
2549:
2548:
2530:
2506:
2500:
2499:
2489:
2465:
2459:
2458:
2422:
2416:
2415:
2413:
2412:
2401:
2395:
2394:
2384:
2366:
2357:
2348:
2347:
2337:
2305:
2299:
2298:
2288:
2256:
2250:
2249:
2239:
2229:
2197:
2191:
2190:
2162:
2156:
2155:
2119:
2113:
2112:
2084:
2075:
2074:
2064:
2049:Pigment Cell Res
2040:
2034:
2033:
1994:Ann N Y Acad Sci
1989:
1983:
1982:
1954:
1945:
1944:
1921:Pigment Cell Res
1916:
1910:
1909:
1901:
1895:
1894:
1884:
1852:
1846:
1845:
1809:
1803:
1802:
1774:
1768:
1767:
1757:
1725:
1719:
1718:
1682:
1676:
1675:
1642:(4295): 1225–7.
1631:
1625:
1624:
1596:
1590:
1589:
1556:(4116): 1034–5.
1545:
1539:
1538:
1515:Pigment Cell Res
1510:
1504:
1503:
1480:Pigment Cell Res
1475:
1462:
1461:
1438:Pigment Cell Res
1433:
1427:
1426:
1398:
1392:
1374:
1368:
1367:
1357:
1321:
1315:
1301:
1295:
1294:
1260:
1251:
1250:
1239:
1233:
1232:
1220:
1214:
1186:
1180:
1179:
1168:10.1670/09-026.1
1147:
1023:biological assay
1013:suppression and
815:molecular motors
811:protein kinase A
807:second messenger
635:, generates its
608:Papua New Guinea
530:At the bottom a
516:metallic colours
352:
351:
345:
334:
333:
327:
320:
319:
313:
281:veiled chameleon
243:
155:
21:
3951:
3950:
3946:
3945:
3944:
3942:
3941:
3940:
3916:
3915:
3914:
3909:
3881:
3866:Nervous system
3841:
3820:
3811:Olfactory organ
3784:
3748:
3693:
3683:
3623:
3582:
3573:
3567:
3561:
3544:
3498:
3431:
3402:Belemnoid guard
3390:
3379:Nautiloid shell
3341:
3335:
3305:
3300:
3229:
3141:
3113:
3031:
3022:
2962:
2959:
2910:
2904:
2900:
2897:
2892:
2871:(2–3): 141–56.
2862:
2861:
2857:
2820:Current Biology
2813:
2812:
2808:
2770:
2769:
2765:
2749:
2745:
2720:
2719:
2715:
2679:
2678:
2674:
2644:
2643:
2639:
2608:
2607:
2603:
2557:
2556:
2552:
2508:
2507:
2503:
2467:
2466:
2462:
2424:
2423:
2419:
2410:
2408:
2403:
2402:
2398:
2364:
2359:
2358:
2351:
2320:(20): 1877–81.
2314:Current Biology
2307:
2306:
2302:
2265:Current Biology
2258:
2257:
2253:
2212:(36): 13204–9.
2199:
2198:
2194:
2164:
2163:
2159:
2121:
2120:
2116:
2086:
2085:
2078:
2042:
2041:
2037:
1991:
1990:
1986:
1956:
1955:
1948:
1918:
1917:
1913:
1903:
1902:
1898:
1854:
1853:
1849:
1811:
1810:
1806:
1776:
1775:
1771:
1727:
1726:
1722:
1684:
1683:
1679:
1633:
1632:
1628:
1598:
1597:
1593:
1547:
1546:
1542:
1512:
1511:
1507:
1477:
1476:
1465:
1435:
1434:
1430:
1400:
1399:
1395:
1375:
1371:
1323:
1322:
1318:
1302:
1298:
1283:
1262:
1261:
1254:
1243:Darwin, Charles
1241:
1240:
1236:
1222:
1221:
1217:
1209:Wayback Machine
1187:
1183:
1152:Boa constrictor
1149:
1148:
1144:
1140:
1118:
1055:
1035:pertussis toxin
972:
873:
852:
831:
691:
671:, and atypical
646:
570:carboxylic acid
566:dihydroxyindole
552:, that appears
542:
524:
504:tapetum lucidum
465:
421:
413:red blood cells
273:
203:
201:Human discovery
183:cell signalling
145:
35:
28:
23:
22:
15:
12:
11:
5:
3949:
3947:
3939:
3938:
3933:
3928:
3918:
3917:
3911:
3910:
3886:
3883:
3882:
3880:
3879:
3878:
3877:
3872:
3864:
3859:
3857:Chromatophores
3853:
3851:
3847:
3846:
3843:
3842:
3840:
3839:
3834:
3828:
3826:
3822:
3821:
3819:
3818:
3813:
3808:
3803:
3798:
3792:
3790:
3786:
3785:
3783:
3782:
3777:
3772:
3767:
3762:
3756:
3754:
3750:
3749:
3747:
3746:
3741:
3740:
3739:
3734:
3729:
3719:
3718:
3717:
3706:
3704:
3703:Brachial crown
3697:
3689:
3688:
3685:
3684:
3682:
3681:
3676:
3671:
3666:
3661:
3656:
3647:
3642:
3637:
3631:
3629:
3625:
3624:
3622:
3621:
3616:
3611:
3606:
3601:
3595:
3593:
3586:
3575:
3574:
3555:
3553:
3550:
3549:
3546:
3545:
3543:
3542:
3537:
3532:
3527:
3522:
3517:
3512:
3506:
3504:
3500:
3499:
3497:
3496:
3491:
3486:
3481:
3476:
3471:
3466:
3461:
3456:
3450:
3448:
3441:
3437:
3436:
3433:
3432:
3430:
3429:
3424:
3419:
3414:
3409:
3404:
3398:
3396:
3392:
3391:
3389:
3388:
3387:
3386:
3376:
3371:
3369:Ammonoid shell
3365:
3363:
3356:
3349:
3343:
3342:
3336:
3334:
3333:
3326:
3319:
3311:
3302:
3301:
3299:
3298:
3293:
3288:
3283:
3278:
3273:
3268:
3263:
3258:
3253:
3248:
3243:
3237:
3235:
3234:Related topics
3231:
3230:
3228:
3227:
3222:
3217:
3212:
3207:
3206:
3205:
3195:
3190:
3185:
3183:Countershading
3180:
3175:
3170:
3165:
3160:
3155:
3149:
3147:
3143:
3142:
3140:
3139:
3134:
3129:
3123:
3121:
3115:
3114:
3112:
3111:
3106:
3101:
3099:Holochroal eye
3096:
3095:
3094:
3089:
3079:
3078:
3077:
3067:
3062:
3057:
3052:
3047:
3041:
3039:
3033:
3032:
3025:
3023:
3021:
3020:
3019:
3018:
3013:
3008:
2998:
2993:
2988:
2983:
2978:
2972:
2970:
2964:
2963:
2960:
2958:
2957:
2950:
2943:
2935:
2929:
2928:
2923:
2918:
2913:
2911:(1.20 MB)
2896:
2895:External links
2893:
2891:
2890:
2855:
2806:
2763:
2743:
2713:
2672:
2637:
2601:
2550:
2501:
2460:
2433:(6): 504–513.
2417:
2396:
2349:
2300:
2251:
2192:
2157:
2114:
2076:
2035:
1984:
1965:(3): 206–213.
1946:
1911:
1896:
1867:(3): 297–301.
1847:
1820:(10): 1581–9.
1804:
1769:
1720:
1677:
1626:
1607:(5): 510–516.
1591:
1540:
1505:
1463:
1428:
1393:
1369:
1340:(3): 493–504.
1316:
1296:
1281:
1252:
1234:
1215:
1181:
1162:(4): 506–515.
1141:
1139:
1136:
1135:
1134:
1129:
1124:
1117:
1114:
1054:
1051:
971:
968:
936:chromatoblasts
872:
869:
830:
827:
823:microfilaments
752:, through its
690:
687:
645:
642:
594:, a pteridine
587:Anolis grahami
523:
520:
464:
461:
420:
417:
272:
269:
249:Charles Darwin
202:
199:
195:drug discovery
49:Chromatophores
26:
24:
18:Chromatophores
14:
13:
10:
9:
6:
4:
3:
2:
3948:
3937:
3936:Pigment cells
3934:
3932:
3929:
3927:
3924:
3923:
3921:
3908:
3904:
3900:
3896:
3892:
3889:
3884:
3876:
3873:
3871:
3868:
3867:
3865:
3863:
3860:
3858:
3855:
3854:
3852:
3848:
3838:
3835:
3833:
3830:
3829:
3827:
3823:
3817:
3814:
3812:
3809:
3807:
3804:
3802:
3799:
3797:
3794:
3793:
3791:
3787:
3781:
3778:
3776:
3773:
3771:
3768:
3766:
3763:
3761:
3758:
3757:
3755:
3753:Buccal region
3751:
3745:
3742:
3738:
3735:
3733:
3730:
3728:
3725:
3724:
3723:
3720:
3716:
3713:
3712:
3711:
3708:
3707:
3705:
3701:
3698:
3696:
3690:
3680:
3677:
3675:
3672:
3670:
3667:
3665:
3662:
3660:
3659:Needham's sac
3657:
3655:
3651:
3648:
3646:
3643:
3641:
3638:
3636:
3633:
3632:
3630:
3626:
3620:
3617:
3615:
3612:
3610:
3607:
3605:
3602:
3600:
3597:
3596:
3594:
3590:
3587:
3585:
3580:
3576:
3571:
3565:
3559:
3541:
3538:
3536:
3533:
3531:
3528:
3526:
3523:
3521:
3518:
3516:
3513:
3511:
3508:
3507:
3505:
3501:
3495:
3492:
3490:
3487:
3485:
3482:
3480:
3477:
3475:
3472:
3470:
3467:
3465:
3462:
3460:
3457:
3455:
3452:
3451:
3449:
3445:
3442:
3438:
3428:
3425:
3423:
3422:Spirula shell
3420:
3418:
3417:Gladius (pen)
3415:
3413:
3410:
3408:
3407:Cirrate shell
3405:
3403:
3400:
3399:
3397:
3393:
3385:
3382:
3381:
3380:
3377:
3375:
3372:
3370:
3367:
3366:
3364:
3360:
3357:
3353:
3350:
3348:
3344:
3339:
3332:
3327:
3325:
3320:
3318:
3313:
3312:
3309:
3297:
3294:
3292:
3289:
3287:
3284:
3282:
3279:
3277:
3274:
3272:
3269:
3267:
3264:
3262:
3259:
3257:
3254:
3252:
3249:
3247:
3244:
3242:
3241:Animal senses
3239:
3238:
3236:
3232:
3226:
3223:
3221:
3218:
3216:
3213:
3211:
3208:
3204:
3201:
3200:
3199:
3196:
3194:
3191:
3189:
3186:
3184:
3181:
3179:
3176:
3174:
3173:Chromatophore
3171:
3169:
3166:
3164:
3161:
3159:
3156:
3154:
3151:
3150:
3148:
3144:
3138:
3135:
3133:
3130:
3128:
3125:
3124:
3122:
3120:
3116:
3110:
3107:
3105:
3102:
3100:
3097:
3093:
3090:
3088:
3085:
3084:
3083:
3080:
3076:
3073:
3072:
3071:
3070:Mammalian eye
3068:
3066:
3063:
3061:
3058:
3056:
3053:
3051:
3048:
3046:
3045:Arthropod eye
3043:
3042:
3040:
3038:
3034:
3029:
3017:
3014:
3012:
3009:
3007:
3004:
3003:
3002:
2999:
2997:
2994:
2992:
2989:
2987:
2984:
2982:
2979:
2977:
2974:
2973:
2971:
2969:
2965:
2956:
2951:
2949:
2944:
2942:
2937:
2936:
2933:
2927:
2924:
2922:
2919:
2917:
2914:
2903:
2899:
2898:
2894:
2886:
2882:
2878:
2874:
2870:
2866:
2859:
2856:
2851:
2847:
2842:
2837:
2833:
2829:
2825:
2821:
2817:
2810:
2807:
2802:
2798:
2794:
2790:
2786:
2782:
2779:(2): 250–80.
2778:
2774:
2767:
2764:
2760:
2756:
2753:
2747:
2744:
2740:
2735:
2731:
2729:
2724:
2717:
2714:
2709:
2705:
2700:
2695:
2692:(2): 169–75.
2691:
2687:
2683:
2676:
2673:
2668:
2664:
2660:
2656:
2653:(1): 111–20.
2652:
2648:
2641:
2638:
2633:
2629:
2625:
2621:
2618:(4): 189–97.
2617:
2613:
2605:
2602:
2597:
2593:
2589:
2585:
2581:
2577:
2573:
2569:
2565:
2561:
2554:
2551:
2546:
2542:
2538:
2534:
2529:
2524:
2521:(4): 326–36.
2520:
2516:
2512:
2505:
2502:
2497:
2493:
2488:
2483:
2480:(2): 277–93.
2479:
2475:
2471:
2464:
2461:
2456:
2452:
2448:
2444:
2440:
2436:
2432:
2428:
2421:
2418:
2406:
2400:
2397:
2392:
2388:
2383:
2378:
2375:(1): 148–60.
2374:
2370:
2363:
2356:
2354:
2350:
2345:
2341:
2336:
2331:
2327:
2323:
2319:
2315:
2311:
2304:
2301:
2296:
2292:
2287:
2282:
2278:
2274:
2270:
2266:
2262:
2255:
2252:
2247:
2243:
2238:
2233:
2228:
2223:
2219:
2215:
2211:
2207:
2203:
2196:
2193:
2188:
2184:
2180:
2176:
2173:(4): 289–95.
2172:
2168:
2161:
2158:
2153:
2149:
2145:
2141:
2137:
2133:
2130:(3): 328–30.
2129:
2125:
2118:
2115:
2110:
2106:
2102:
2098:
2095:(2): 184–91.
2094:
2090:
2083:
2081:
2077:
2072:
2068:
2063:
2058:
2055:(5): 454–60.
2054:
2050:
2046:
2039:
2036:
2031:
2027:
2023:
2019:
2015:
2011:
2007:
2003:
2000:(1): 319–30.
1999:
1995:
1988:
1985:
1980:
1976:
1972:
1968:
1964:
1960:
1953:
1951:
1947:
1942:
1938:
1934:
1930:
1926:
1922:
1915:
1912:
1907:
1900:
1897:
1892:
1888:
1883:
1878:
1874:
1870:
1866:
1862:
1858:
1851:
1848:
1843:
1839:
1835:
1831:
1827:
1823:
1819:
1815:
1808:
1805:
1800:
1796:
1792:
1788:
1784:
1780:
1773:
1770:
1765:
1761:
1756:
1751:
1747:
1743:
1739:
1735:
1731:
1724:
1721:
1716:
1712:
1708:
1704:
1700:
1696:
1692:
1688:
1681:
1678:
1673:
1669:
1665:
1661:
1657:
1653:
1649:
1645:
1641:
1637:
1630:
1627:
1622:
1618:
1614:
1610:
1606:
1602:
1595:
1592:
1587:
1583:
1579:
1575:
1571:
1567:
1563:
1559:
1555:
1551:
1544:
1541:
1536:
1532:
1528:
1524:
1521:(5): 523–31.
1520:
1516:
1509:
1506:
1501:
1497:
1493:
1489:
1486:(5): 300–19.
1485:
1481:
1474:
1472:
1470:
1468:
1464:
1459:
1455:
1451:
1447:
1443:
1439:
1432:
1429:
1424:
1420:
1416:
1412:
1409:(3): 405–16.
1408:
1404:
1397:
1394:
1391:
1390:0-19-509861-7
1387:
1383:
1379:
1373:
1370:
1365:
1361:
1356:
1351:
1347:
1343:
1339:
1335:
1331:
1329:
1320:
1317:
1314:
1313:0-520-02347-1
1310:
1306:
1300:
1297:
1292:
1288:
1284:
1278:
1274:
1270:
1266:
1259:
1257:
1253:
1248:
1244:
1238:
1235:
1230:
1226:
1219:
1216:
1213:
1210:
1206:
1203:
1199:
1195:
1191:
1185:
1182:
1177:
1173:
1169:
1165:
1161:
1157:
1153:
1146:
1143:
1137:
1133:
1130:
1128:
1125:
1123:
1120:
1119:
1115:
1113:
1111:
1107:
1103:
1098:
1093:
1091:
1087:
1083:
1079:
1075:
1071:
1064:
1059:
1052:
1050:
1048:
1044:
1040:
1036:
1032:
1028:
1024:
1020:
1016:
1012:
1007:
1002:
1000:
996:
995:
990:
986:
982:
978:
969:
967:
965:
961:
960:
955:
954:
949:
945:
941:
940:fertilization
937:
933:
930:When and how
928:
926:
922:
917:
913:
909:
905:
901:
897:
893:
885:
881:
880:Cross-section
877:
870:
868:
866:
861:
857:
851:
843:
839:
835:
828:
826:
824:
820:
816:
812:
808:
804:
800:
796:
792:
788:
787:
782:
777:
775:
771:
767:
763:
759:
758:melanocortins
755:
751:
750:noradrenaline
746:
742:
738:
730:
726:
721:
717:
713:
711:
707:
706:colour change
705:
704:physiological
695:
688:
686:
684:
683:
678:
674:
670:
666:
661:
657:
656:
651:
643:
638:
634:
633:
627:
623:
621:
617:
611:
609:
605:
602:species from
601:
597:
593:
589:
588:
583:
579:
575:
571:
567:
563:
559:
555:
551:
547:
541:
533:
528:
521:
519:
517:
513:
509:
505:
500:
498:
494:
490:
486:
482:
481:Bragg mirrors
478:
469:
462:
460:
458:
454:
450:
446:
441:
438:
434:
430:
426:
418:
416:
414:
410:
406:
402:
397:
395:
390:
386:
382:
378:
374:
370:
365:
363:
358:
356:
346:
344:
338:
328:
326:
314:
312:
306:
302:
298:
294:
293:chromatophore
286:
282:
277:
270:
267:
262:
260:
259:
254:
250:
246:
244:
242:
236:
232:
227:
223:
221:
220:
215:
211:
207:
200:
198:
196:
192:
188:
184:
180:
176:
172:
168:
164:
143:
141:
137:
133:
129:
125:
121:
117:
116:erythrophores
113:
109:
105:
101:
97:
92:
90:
86:
82:
78:
74:
70:
66:
62:
58:
54:
50:
44:
39:
33:
19:
3887:
3856:
3816:Nuchal organ
3715:Hectocotylus
3604:Fins (wings)
3510:Body chamber
3474:Periostracum
3172:
3104:Parietal eye
3050:Compound eye
2868:
2864:
2858:
2823:
2819:
2809:
2776:
2772:
2766:
2754:
2746:
2737:
2734:the original
2726:
2716:
2689:
2685:
2675:
2650:
2646:
2640:
2615:
2611:
2604:
2563:
2559:
2553:
2518:
2514:
2504:
2477:
2473:
2463:
2430:
2426:
2420:
2409:. Retrieved
2407:. 2023-06-13
2399:
2372:
2368:
2317:
2313:
2303:
2271:(3): 165–8.
2268:
2264:
2254:
2209:
2205:
2195:
2170:
2166:
2160:
2127:
2123:
2117:
2092:
2088:
2052:
2048:
2038:
1997:
1993:
1987:
1962:
1958:
1927:(1): 59–64.
1924:
1920:
1914:
1905:
1899:
1864:
1860:
1850:
1817:
1813:
1807:
1782:
1778:
1772:
1740:(1): 67–79.
1737:
1733:
1723:
1693:(4): 614–7.
1690:
1686:
1680:
1639:
1635:
1629:
1604:
1600:
1594:
1553:
1549:
1543:
1518:
1514:
1508:
1483:
1479:
1444:(1): 28–36.
1441:
1437:
1431:
1406:
1402:
1396:
1377:
1372:
1337:
1333:
1327:
1319:
1304:
1299:
1264:
1246:
1237:
1228:
1224:
1218:
1212:Google books
1197:
1189:
1184:
1159:
1155:
1151:
1145:
1094:
1086:translucency
1068:
1003:
992:
988:
973:
957:
951:
947:
935:
929:
912:basal lamina
896:neural crest
889:
855:
853:
819:microtubules
798:
784:
778:
734:
714:
702:
700:
680:
676:
653:
647:
630:
620:phaeomelanin
612:
585:
548:, a type of
543:
522:Melanophores
501:
474:
455:-restricted
442:
422:
398:
366:
362:biochemistry
359:
340:
337:chromatocyte
336:
322:
308:
296:
292:
290:
284:
264:
256:
247:
238:
235:invertebrate
229:
225:
217:
204:
170:
166:
165:, is called
160:
139:
136:melanophores
135:
131:
119:
115:
112:xanthophores
111:
104:neural crest
93:
48:
47:
3903:Egg fossils
3862:Photophores
3825:Other parts
3770:Odontophore
3692:Head &
3525:Phragmocone
3281:Pseudopupil
3163:Aposematism
3082:Mollusc eye
2369:J Neurobiol
1785:(1): 9–20.
1734:J Cell Biol
1334:J Cell Biol
1188:Aristotle.
1127:Chromophore
999:skin colour
932:multipotent
900:neural tube
871:Development
795:hair colour
673:dichromatic
669:glass frogs
644:Cyanophores
616:melanocytes
592:pterorhodin
449:metabolised
394:iridescence
389:wavelengths
377:carotenoids
301:vertebrates
171:metachrosis
140:cyanophores
132:leucophores
120:iridophores
100:ectothermic
85:melanocytes
73:cephalopods
69:crustaceans
3920:Categories
3837:Statocysts
3674:Osphradium
3412:Cuttlebone
3338:Cephalopod
3271:Ommatidium
3203:coincident
3168:Camouflage
3146:Coloration
3087:cephalopod
2981:Chameleons
2755:PhD Thesis
2411:2024-06-08
1097:cuttlefish
1078:cuttlefish
1063:cuttlefish
1061:An infant
1047:cuttlefish
1031:biosensors
966:disorder.
850:Camouflage
848:See also:
729:time-lapse
578:tyrosinase
572:with some
540:Melanocyte
538:See also:
381:pteridines
369:biochromes
355:melanocyte
297:chromoforo
253:cuttlefish
214:camouflage
179:chameleons
163:camouflage
128:iridescent
124:reflective
114:(yellow),
96:eye colour
89:coloration
57:amphibians
3895:Paralarva
3722:Tentacles
3535:Siphuncle
3494:Umbilicus
3479:Sculpture
3384:Orthocone
3286:Rhopalium
3119:Evolution
3092:gastropod
3060:Eye shine
3055:Eagle eye
2986:Dinosaurs
2739:uniforms.
2167:Zebrafish
2124:Nat Genet
1303:Fox, DL.
1132:Thylakoid
1006:biomarker
964:leucistic
921:optic cup
904:caldesmon
838:Zebrafish
813:to drive
781:homologue
770:paracrine
762:melatonin
725:zebrafish
723:A single
604:Australia
600:tree frog
546:eumelanin
499:colours.
291:The term
241:cromoforo
206:Aristotle
134:(white),
3760:Aptychus
3732:Dactylus
3503:Internal
3454:Aperture
3447:External
3440:Features
3395:Internal
3362:External
3153:Albinism
2850:25042589
2801:26566732
2757:. 2005;
2667:15967358
2632:11033434
2588:16357253
2545:15112204
2537:15250934
2496:10985850
2474:Dev Biol
2455:30194336
2447:12242708
2391:12486702
2344:15498498
2246:15331778
2187:19133827
2109:12620396
2089:Genomics
2071:15357831
2030:37853710
2022:12851332
1979:16704454
1941:12519126
1891:12551954
1834:14706537
1814:Peptides
1707:21501419
1672:45014819
1621:12950730
1586:33900017
1535:12950732
1500:11041206
1384:, 1998.
1205:Archived
1202:Abstract
1176:53634890
1116:See also
1039:military
1011:appetite
985:melanoma
946:such as
908:ectoderm
754:receptor
745:neuronal
741:hormonal
562:tyrosine
556:or dark-
457:crickets
453:carotene
437:vesicles
405:albinism
403:, as in
401:mutation
261:(1860):
187:hormones
65:reptiles
3850:General
3650:Ink sac
3515:Camerae
3489:Sutures
3340:anatomy
3215:Mimicry
3188:Crypsis
3001:Mammals
2885:1422807
2828:Bibcode
2793:5700268
2750:Lee I.
2708:1936946
2596:2245002
2568:Bibcode
2560:Science
2322:Bibcode
2295:9443917
2273:Bibcode
2214:Bibcode
2152:7980311
2144:7581459
2002:Bibcode
1882:2172679
1842:2593619
1799:2543509
1764:5691979
1755:2107474
1715:3299089
1644:Bibcode
1636:Science
1578:4748673
1558:Bibcode
1550:Science
1458:7792252
1423:5769930
1364:5885426
1355:2106771
1291:5337298
1231:: 1–13.
1110:texture
1090:opacity
1070:Coleoid
1015:tanning
994:Slc24a5
916:somites
737:teleost
574:pyrrole
550:melanin
508:purines
485:Tyndall
477:guanine
210:octopus
175:octopus
118:(red),
106:during
77:Mammals
53:pigment
3775:Radula
3727:Carpus
3584:funnel
3581:&
3579:Mantle
3540:Whorls
3464:Callus
3427:Stylet
3006:horses
2968:Vision
2909:
2883:
2848:
2799:
2791:
2706:
2665:
2630:
2594:
2586:
2543:
2535:
2494:
2453:
2445:
2389:
2342:
2293:
2244:
2237:516548
2234:
2185:
2150:
2142:
2107:
2069:
2028:
2020:
1977:
1939:
1889:
1879:
1840:
1832:
1797:
1762:
1752:
1713:
1705:
1670:
1664:860137
1662:
1619:
1584:
1576:
1533:
1498:
1456:
1421:
1388:
1362:
1352:
1311:
1289:
1279:
1174:
1074:squids
989:golden
956:, and
925:retina
865:anoles
764:, and
710:dermis
637:violet
532:mutant
433:orange
425:yellow
325:phoros
311:chrōma
307:words
3926:Cells
3891:Spawn
3737:Manus
3695:limbs
3530:Septa
3520:Nacre
3484:Spire
3469:Lirae
3355:Types
3347:Shell
3075:human
2996:Toads
2976:Birds
2905:(PDF)
2797:S2CID
2592:S2CID
2541:S2CID
2451:S2CID
2365:(PDF)
2148:S2CID
2026:S2CID
1838:S2CID
1711:S2CID
1668:S2CID
1582:S2CID
1172:S2CID
1138:Notes
1082:glial
1019:drugs
977:model
953:sox10
884:trunk
860:mimic
596:dimer
558:brown
554:black
512:white
497:green
350:κύτος
343:kytos
332:φόρος
318:χρῶμα
305:Greek
81:birds
43:squid
3832:Eyes
3765:Beak
3710:Arms
3652:and
3619:Tail
3459:Apex
3037:Eyes
3016:cats
3011:dogs
2991:Fish
2881:PMID
2846:PMID
2789:PMID
2730:News
2704:PMID
2663:PMID
2628:PMID
2584:PMID
2533:PMID
2492:PMID
2443:PMID
2387:PMID
2340:PMID
2291:PMID
2242:PMID
2183:PMID
2140:PMID
2105:PMID
2067:PMID
2018:PMID
1975:PMID
1937:PMID
1887:PMID
1830:PMID
1795:PMID
1760:PMID
1703:PMID
1660:PMID
1617:PMID
1574:PMID
1531:PMID
1496:PMID
1454:PMID
1419:PMID
1386:ISBN
1360:PMID
1309:ISBN
1287:PMID
1277:ISBN
1154:)".
1076:and
959:mitf
821:and
799:MC1R
793:and
791:skin
786:MC1R
667:and
660:cyan
606:and
495:or -
493:blue
409:Haem
379:and
371:and
87:for
79:and
71:and
61:fish
3654:ink
2873:doi
2836:doi
2781:doi
2694:doi
2655:doi
2620:doi
2576:doi
2564:310
2523:doi
2482:doi
2478:225
2435:doi
2377:doi
2330:doi
2281:doi
2232:PMC
2222:doi
2210:101
2175:doi
2132:doi
2097:doi
2057:doi
2010:doi
1998:994
1967:doi
1929:doi
1877:PMC
1869:doi
1865:160
1822:doi
1787:doi
1750:PMC
1742:doi
1695:doi
1652:doi
1640:196
1609:doi
1566:doi
1554:182
1523:doi
1488:doi
1446:doi
1411:doi
1350:PMC
1342:doi
1269:doi
1164:doi
948:kit
783:of
743:or
487:or
429:red
255:in
189:or
169:or
130:),
98:in
3922::
3905:•
3893:→
2879:.
2869:40
2867:.
2844:.
2834:.
2824:24
2822:.
2818:.
2795:.
2787:.
2777:89
2775:.
2725:.
2702:.
2690:66
2688:.
2684:.
2661:.
2651:21
2649:.
2626:.
2616:42
2614:.
2590:.
2582:.
2574:.
2562:.
2539:.
2531:.
2519:17
2517:.
2513:.
2490:.
2476:.
2472:.
2449:.
2441:.
2431:58
2429:.
2385:.
2373:54
2371:.
2367:.
2352:^
2338:.
2328:.
2318:14
2316:.
2312:.
2289:.
2279:.
2267:.
2263:.
2240:.
2230:.
2220:.
2208:.
2204:.
2181:.
2169:.
2146:.
2138:.
2128:11
2126:.
2103:.
2093:81
2091:.
2079:^
2065:.
2053:17
2051:.
2047:.
2024:.
2016:.
2008:.
1996:.
1973:.
1963:19
1961:.
1949:^
1935:.
1925:16
1923:.
1885:.
1875:.
1863:.
1859:.
1836:.
1828:.
1818:24
1816:.
1793:.
1783:13
1781:.
1758:.
1748:.
1738:38
1736:.
1732:.
1709:.
1701:.
1691:24
1689:.
1666:.
1658:.
1650:.
1638:.
1615:.
1605:16
1603:.
1580:.
1572:.
1564:.
1552:.
1529:.
1519:16
1517:.
1494:.
1484:13
1482:.
1466:^
1452:.
1440:.
1417:.
1407:12
1405:.
1380:,
1358:.
1348:.
1338:27
1336:.
1332:.
1285:.
1275:.
1255:^
1227:.
1170:.
1160:44
1158:.
1001:.
950:,
927:.
825:.
760:,
685:.
622:.
283:,
279:A
197:.
126:/
91:.
75:.
67:,
63:,
59:,
3897:(
3330:e
3323:t
3316:v
2954:e
2947:t
2940:v
2907:.
2887:.
2875::
2852:.
2838::
2830::
2803:.
2783::
2761:.
2710:.
2696::
2669:.
2657::
2634:.
2622::
2598:.
2578::
2570::
2547:.
2525::
2498:.
2484::
2457:.
2437::
2414:.
2393:.
2379::
2346:.
2332::
2324::
2297:.
2283::
2275::
2269:8
2248:.
2224::
2216::
2189:.
2177::
2171:5
2154:.
2134::
2111:.
2099::
2073:.
2059::
2032:.
2012::
2004::
1981:.
1969::
1943:.
1931::
1893:.
1871::
1844:.
1824::
1801:.
1789::
1766:.
1744::
1717:.
1697::
1674:.
1654::
1646::
1623:.
1611::
1588:.
1568::
1560::
1537:.
1525::
1502:.
1490::
1460:.
1448::
1442:8
1425:.
1413::
1366:.
1344::
1293:.
1271::
1229:9
1178:.
1166::
431:/
347:(
339:(
329:(
315:(
122:(
34:.
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
