610:(the squinting bush brown butterfly) have shown that overexpression or down-regulation of Dll in the first expression domain correlates with bigger and smaller eyespots respectively. However, if this is done on the second domain then the overall size of the eyespots remains the same, but the width of the black ring raises with a higher amount of Dll. This suggests that Dll might be responsible for the differentiation of the focus in the first expression domain and might be involved in establishing the ring colour patterns in the second domain. These experiments together with the wide distribution of Dll across eyespot forming butterflies suggest that this transcription factor is a central regulator for the correct patterning of the eyespots.
401:
359:
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382:
116:
319:
126:
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2177:
694:, forest users wear face masks with eye markings on the backs of their heads in the hope of reducing tiger attacks. In the study on 2061 cattle in 14 herds over 4 years, 683 were given eye markings, 543 were painted with crosses, and 835 were unpainted. None of the eyed cattle were predated, but 4 cross-marked and 15 unmarked cattle were killed, one by a leopard, the rest by lions. Both the eyespots and the cross markings provided
256:
472:
22:
683:
588:
The Distal-less gene is present in almost all eyespot organizers, making it an ideal candidate to carry out major functions of eyespot formation. During the wing imaginal disc development Dll, has two expression domains separated by a temporal component. First Dll is expressed in a group of cells in
503:
model, but could be explained by either a source/threshold model, in which the focus creates the morphogen, or by the sink model, in which the focus generates a gradient by removing a morphogen which was created elsewhere. Several genes involved in eyespot formation have been identified that can fit
631:
fruit flies, Hh is expressed in all cells in the posterior compartment of the developing butterfly wing during the mid fifth instar of butterfly wing development. However, in butterflies, Hh expression is significantly higher in those cells that flank the potential foci. Higher transcription levels
635:
Furthermore, cells that are flanked by the cells expressing the highest level of Hh signaling are fated to become the foci, indicating that focus cell fate determination relies on high concentrations of Hh in surrounding cells. However, this observation has not been totally confirmed as a rule for
624:
The
Hedgehog (Hh) gene is the other element that has been functionally tested in the formation of eyespots. Investigating genes involved in wing development and morphogenetic activity has led to the discovery that Hh has a primary role in the morphogenetic signaling center of the foci. In a manner
658:
The Notch (N) gene expression precedes an upregulation of Dll in the cells that will become the center of the focus. This makes N the earliest developmental signal, so far studied, that is related with the establishment of the eyespots. Loss of N completely disrupts Dll expression, and eventually
632:
of Hh, along with other known associates of the Hh pathway, namely patched (Ptc) the Hh receptor, and cubitus interruptus (Ci), the Hh transcription factor is seen throughout the mid to late fifth instar as well, which further implies a role for Hh signaling in eyespot development and patterning.
207:
butterflies; they commonly perch upside down with the false head raised and shift their rear wings repeatedly, causing antenna-like movements of the "tails" on their wings. Studies of rear-wing damage support the hypothesis that this deflects attacks from the insect's head.
659:
eyespot formation, in several butterfly species. A variety of other wing patterns are determined by N and Dll patterns of expression in early development of the wing imaginal disc, suggesting that a single mechanism patterns multiple coloration structures of the wing.
64:, to draw a predator's attention away from the prey's most vulnerable body parts. Or they may serve to make the prey appear inedible or dangerous. Eyespot markings may play a role in intraspecies communication or courtship; a well-known example is the eyespots on a
667:
Butterfly eyespots are formed by an interplay of at least 3 genes, namely Distal-less (Dll), spalt (sal), and
Antennapedia (Antp), hence their evolution has been shaped by differential expression of these genes in different butterfly taxa, as shown in
527:
which patterns the wings of other insects. This rogue regulatory circuit is able to pattern both the anterior and posterior eyespots independent of the usual anterior/posterior wing compartmentalization restrictions seen in the fruit fly
173:; this is a response to a seasonal fall in temperature, causing a shift in selection towards smaller, less conspicuous eyespots among those individuals developing at that time. Butterfly eyespots may play a role in mate recognition and
188:), have eyespots on their anterior abdominal segments. When alarmed, they retract the head and the thoracic segments into the body, leaving the apparently threatening large eyes at the front of the visible part of the body.
944:
498:
on the butterfly wing eyespot foci to demonstrate that a long range signaling mechanism or morphogen gradient controlled the formation of eyespots in both space and time. The findings cannot be explained by a simple
1929:
Murugesan, Suriya
Narayanan; Connahs, Heidi; Matsuoka, Yuji; Gupta, Mainak Das; Tiong, Galen J. L.; Huq, Manizah; Gowri, V.; Monroe, Sarah; Deem, Kevin D.; Werner, Thomas; Tomoyasu, Yoshinori (2022-02-22).
600:
stage. The second domain starts around 20 hours after pupation around the original central cluster of cells, in an area in which a black ring of the eyespot will be formed. Functional experiments using
312:, maintain their eyespots throughout their adult lives. These eyespots can take a form very similar to those seen in most butterflies, with a focus surrounded by concentric rings of other pigmentation.
1673:
Breakfield, M.P.; Gates, J.; Keys, D.; Kesbeke, F.; Wijngaarden, J. P.; Monteiro, A.; French, V.; Carroll, S. B. (November 1996). "Development, plasticity and evolution of butterfly eyespot patterns".
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338:
296:
gets its name from a large and conspicuous eyespot on each side of the body near the tail. A black vertical bar on the head runs through the true eye, making it hard to see. This may deceive
686:
Eyespot experiment on cattle in
Botswana. Both the eyespots (left) and the cross markings (centre) protected the cattle from predation by lions, compared to the unmarked controls (right).
163:, the conspicuous eyespots are hidden at rest to decrease detectability, and only exposed when they believe potential predators are nearby. Butterfly eyespots can mimic dead leaves for
640:. Here they observed that both seem to be expressed in eyespots, suggesting a relation with the Hh signaling pathway. However, other studies did not find evidence of Hh expression in
300:
in two ways: into attacking the tail rather than the more vulnerable head, and about the fish's likely direction of travel. The foureye butterflyfish eyespot is thus an example of
787:
358:
400:
698:
protection. The cattle were always in mixed groups of marked and unmarked animals; it is not known whether marking all animals in a herd would provide effective protection.
274:
females. The number of eyespots in a peacock's train predicts his mating success; when a peacock's train is experimentally pruned, females lose interest. Several species of
546:(En), breaking the anterior/posterior compartmentalization restrictions through increased localized levels of Hh signaling. In turn, this raises expression of its receptor
574:
of existing regulatory mechanisms in butterfly eyespot locus development has given more insight into a fundamental mechanism for the evolution of novel structures.
243:, have white markings, whether spots or bars, on the backs of their ears; it is possible that these signal "follow me" to the young of the species. There may be an
878:
Prudic, K. L.; Jeon, C.; Cao, H.; Monteiro, A. (2011-01-06). "Developmental
Plasticity in Sexual Roles of Butterfly Species Drives Mutual Sexual Ornamentation".
318:
690:
Eyespots painted on the rumps of cows have been shown to reduce cattle predation in Africa. The study authors, Cameron
Radford and colleagues, note that in the
177:. Sexual selection drives the diversification of eyespots in different species of butterflies, as mates select for characteristics like size and brightness.
419:
1339:
Graipel, MaurĂcio
Eduardo; Bogoni, Juliano AndrĂ©; Giehl, Eduardo LuĂs Hettwer; Cerezer, Felipe O.; Cáceres, Nilton Carlos; Eizirik, Eduardo (2019).
1208:
Sourakov, Andrei (2013). "Two heads are better than one: false head allows
Calycopis cecrops (Lycaenidae) to escape predation by a Jumping Spider,
736:
Kodandaramaiah, U. (2011). "Eyespot evolution: phylogenetic insights from
Junonia and related butterfly genera (Nymphalidae: Junoniini)".
491:
signalling centre or organizer, called the focus. This induces neighbouring cells to produce specific pigments which pattern the eyespot.
2372:
381:
1803:
French, V.; Breakfield, P. M. (September 1992). "The development of eyespot patterns on butterfly wings: morphogen sources or sinks?".
812:
2268:
1510:
636:
multiple butterfly species. Studies tried to extrapolate the result of Hh pathway involvement by looking for the expression of Ci in
304:. For the same reason, many juvenile fish display eyespots that disappear during their adult phase. Some species of fish, like the
278:
bear false eyes on the back of the head, misleading predators into reacting as though they were the subject of an aggressive stare.
195:) have filamentous "tails" at the ends of their wings and nearby patterns of markings, which combine to create a "false head". This
1480:
1255:
Robbins, Robert K. (November 1981). "The 'False Head' Hypothesis: Predation and Wing
Pattern Variation of Lycaenid Butterflies".
3238:
1932:"Butterfly eyespots evolved via cooption of an ancestral gene-regulatory network that also patterns antennae, legs, and wings"
2556:
1323:
990:
Bhardwaj, Shivam; Jolander, Lim Si-Hui; Wenk, Markus R.; Oliver, Jeffrey C.; Nijhout, H. Frederik; Monteiro, Antonia (2020).
589:
the center of what will become the focus and eventually the eyespot. This expression starts during the middle of the fifth
3203:
2447:
3048:
2776:
2675:
1890:"Butterfly Wing Pattern Evolution Is Associated with Changes in a Notch/Distal-less Temporal Pattern Formation Process"
567:
76:
3262:
2748:
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1743:
Keys, D. N.; Lewis, D. L.; Selegue, J. E.; Pearson, B. J.; Goodrich, L. V.; Johnson, R. L.; Gates, J.; Scott, M. P.;
220:
of Europe, have eyespots; in the sand lizard's case, there is a row of spots along the back, and a row on each side.
3005:
2791:
2786:
2622:
1605:"Eyespot colour pattern determination by serial induction in fish: Mechanistic convergence with butterfly eyespots"
2365:
3182:
1161:"Male Bicyclus anynana butterflies choose females on the basis of their ventral UV-reflective eyespot centers"
831:"Prey survival by predator intimidation: an experimental study of peacock butterfly defence against blue tits"
3033:
2300:
149:; indeed, it is unclear whether they actually mimic eyes. There is evidence that eyespots in butterflies are
3000:
2484:
2176:
1341:"Melanism evolution in the cat family is influenced by intraspecific communication under low visibility"
3192:
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779:
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in which a spot on the body of an animal resembles an eye of a different animal, to deceive potential
2885:
2551:
695:
534:. The altered regulatory circuit redeploys early developmental signaling sources, like the canonical
2229:
1406:
Petrie, Marion; Halliday, T.; Sanders, C. (1991). "Peahens prefer peacocks with elaborate trains".
1998:
Radford, Cameron; McNutt, John Weldon; Rogers, Tracey; Maslen, Ben; Jordan, Neil (7 August 2020).
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1747:(January 1999). "Recruitment of a hedgehog Regulatory Circuit in Butterfly Eyespot Evolution".
945:"The role of eyespots as anti-predator mechanisms, principally demonstrated in the Lepidoptera"
305:
244:
2193:
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to intimidate predators, or to deflect attacks away from vital body parts. In species such as
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into these models, but only two of them have been functionally tested. These genes are the
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1051:"The use of chemical and visual cues in female choice in the butterfly Bicyclus anynana"
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1554:"Soft corals provide microhabitat for camouflaged juveniles of the Blackspotted wrasse
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1100:"Female choice depends on size but not symmetry of dorsal eyespots in the butterfly
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923:
52:
Eyespots could be explained in at least three different ways. They may be a form of
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992:"Origin of the mechanism of phenotypic plasticity in satyrid butterfly eyespots"
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2156:
2016:
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259:
1906:
1889:
1581:
960:
270:, have conspicuous eyespots in their plumage, used to signal their quality to
75:) of eyespots in a wide variety of animals is controlled by a small number of
49:-like marking. They are found in butterflies, reptiles, cats, birds and fish.
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Artificial eyespots have been shown to reduce predation of cattle by lions.
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1981:
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886:(6013). American Association for the Advancement of Science (AAAS): 73–75.
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130:
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2114:
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1108:
Proceedings of the Royal Society of London. Series B: Biological Sciences
583:
539:
387:
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352:) in a defensive posture, with black eyespots visible just above its head
217:
196:
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138:
84:
61:
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247:
in this case between nocturnal camouflage and intraspecific signalling.
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627:
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552:
530:
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200:
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185:
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1443:"Peacocks with low mating success are more likely to suffer predation"
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2762:
2705:
2538:
2494:
2474:
1694:
1534:
520:
500:
275:
204:
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2000:"Artificial eyespots on cattle reduce predation by large carnivores"
1993:
1991:
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3053:
2572:
1519:
1268:
297:
57:
21:
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of Ci, thereby preventing Ptc expression. From the perspective of
556:, engrailed acts in the posterior compartment to restrict Ptc and
2816:
2634:
2607:
2248:
2083:
1988:
871:
547:
495:
267:
255:
224:
65:
53:
471:
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425:
240:
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1928:
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519:
Butterfly eyespot morphology appears to be the result of the
146:
3094:
597:
2052:
1849:"Origin, Development, and Evolution of Butterfly Eyespots"
1672:
989:
828:
2456:
1997:
1338:
829:
Vallin, A.; Jakobsson, S.; Lind, J.; Wiklund, C. (2005).
46:
199:
misdirects predators such as birds and jumping spiders (
1742:
1055:
Proceedings of the Royal Society B: Biological Sciences
835:
Proceedings of the Royal Society B: Biological Sciences
1305:
1091:
1405:
983:
877:
141:
and certain other insects, as well as birds like the
1603:
Otaki, Joji M.; Ohno, Yoshikazu (28 February 2012).
1332:
1158:
1097:
3249:
2756:
1802:
1311:
1098:Breuker, Casper J.; Brakefield, Paul M. (2002).
1048:
494:Early experiments on eyespot morphogenesis used
18:Eye-like marking used for mimicry or distraction
1936:Proceedings of the National Academy of Sciences
562:(Ci) expression to the anterior compartment by
432:) from back, ocelli clearly visible on the ears
1481:"Northern Pygmy Owl (Glaucidium californicum)"
735:
2742:
2366:
2068:
1668:
1666:
1664:
1318:. University of Chicago Press. pp. 8–9.
935:
933:
778:
550:(Ptc) and transcription factor. Normally, in
1159:Huq, M.; Bhardwaj, S.; Monteiro, A. (2019).
1842:
1840:
1838:
1836:
110:
2749:
2735:
2373:
2359:
2075:
2061:
1887:
1661:
1551:
930:
211:
105:
2269:Coloration evidence for natural selection
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2015:
1971:
1905:
1864:
1824:
1798:
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1786:
1644:
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1135:
1074:
1025:
1007:
854:
806:
475:Plan of a typical butterfly, showing the
1846:
1833:
1483:. Owl Research Institute. Archived from
1207:
681:
470:
266:Male birds of some species, such as the
254:
124:
114:
71:The pattern-forming biological process (
20:
1738:
1736:
1734:
1732:
1730:
1728:
1726:
1724:
1722:
1720:
1541:. Oxford University Press. p. 373.
1505:
1312:Sunquist, Mel; Sunquist, Fiona (2017).
1254:
939:
788:"Do animal eyespots really mimic eyes?"
60:or prey species. They may be a form of
3250:
3239:Index of evolutionary biology articles
1783:
1440:
677:
625:that is similar to the development of
479:foci on the wings that create eyespots
2730:
2557:Evolution of color vision in primates
2380:
2354:
2056:
570:, understanding the redeployment and
512:(a signalling substance that binds a
483:Butterfly eyespots are formed during
203:). Spectacular examples occur in the
77:genes active in embryonic development
1881:
1717:
1533:
1049:Costanzo, K.; Monteiro, A. (2006).
13:
3049:Evolutionary developmental biology
2209:Pouyannian (with pseudocopulation)
1888:Reed, R. D.; Serfas, M.S. (2004).
1866:10.1146/annurev-ento-010814-020942
568:evolutionary developmental biology
332:) displaying eyespots when alarmed
14:
3279:
145:, serve functions in addition to
3006:Evolution of sexual reproduction
2446:
2335:
2334:
2175:
2110:Aristotelian/Distraction display
750:10.1111/j.1525-142X.2009.00357.x
460:
437:
418:
399:
380:
357:
337:
317:
33:), have eyespots on their wings.
1922:
1596:
1545:
1527:
1499:
1473:
1434:
1399:
1283:
1248:
1201:
451:) has false eyes near tail and
191:Butterflies such as the blues (
25:Many butterflies, such as this
2777:Genotype–phenotype distinction
1539:Adaptive Coloration in Animals
1152:
1042:
822:
772:
729:
577:
184:, such as those of hawkmoths (
137:The eye-like markings in some
1:
3034:Regulation of gene expression
2301:Frequency-dependent selection
1420:10.1016/S0003-3472(05)80484-1
1293:. Amphibian and Reptile Group
723:
523:of an altered version of the
292:Some fish have eyespots. The
79:, including the genes called
3204:Endless Forms Most Beautiful
2984:Evolution of genetic systems
2792:Gene–environment correlation
2787:Gene–environment interaction
1769:10.1126/science.283.5401.532
1459:10.1016/0003-3472(92)90072-H
1366:10.1371/journal.pone.0226136
1234:10.1080/00222933.2012.759288
662:
7:
3183:Christiane NĂĽsslein-Volhard
2485:Simple eye in invertebrates
2082:
1853:Annual Review of Entomology
701:
613:
413:) with eyespots behind head
250:
216:Some reptiles, such as the
167:from predators, as seen in
10:
3284:
3059:Hedgehog signaling pathway
2936:Developmental architecture
2681:Infrared sensing in snakes
2017:10.1038/s42003-020-01156-0
1847:Monteiro, Antonia (2015).
1556:Macropharyngodon meleagris
1214:Journal of Natural History
651:
617:
581:
464:
285:
281:
3236:
3215:
3144:
3072:
3026:
3019:
2983:
2935:
2899:
2886:Transgressive segregation
2832:
2769:
2653:
2565:
2552:Evolution of color vision
2537:
2455:
2444:
2386:
2329:
2222:
2184:
2173:
2090:
1907:10.1016/j.cub.2004.06.046
1582:10.1007/s12526-015-0332-x
1552:Bos, A. R. (March 2016).
1165:Journal of Insect Science
961:10.1017/S1464793105006810
696:statistically significant
508:Distalless (Dll) and the
3263:Antipredator adaptations
2230:Anti-predator adaptation
1509:; Pauly, Daniel (eds.).
647:
564:repressing transcription
375:) has false head at rear
151:antipredator adaptations
111:In butterflies and moths
3064:Notch signaling pathway
3039:Gene regulatory network
2922:Dual inheritance theory
1956:10.1073/pnas.2108661119
1257:The American Naturalist
900:10.1126/science.1197114
654:Notch signaling pathway
411:Glaucidium californicum
394:) with rows of eyespots
212:In reptiles and mammals
106:Zoological distribution
97:Notch signaling pathway
3112:cis-regulatory element
3020:Control of development
2900:Non-genetic influences
2866:evolutionary landscape
2004:Communications Biology
1315:Wild Cats of the World
1210:Phidippus pulcherrimus
1120:10.1098/rspb.2002.2005
1067:10.1098/rspb.2006.3729
847:10.1098/rspb.2004.3034
808:10.1093/czoolo/60.1.26
687:
480:
263:
245:evolutionary trade-off
134:
122:
34:
3223:Nature versus nurture
3127:Cell surface receptor
3044:Evo-devo gene toolkit
2943:Developmental biology
2881:Polygenic inheritance
2807:Quantitative genetics
2686:Monocular deprivation
2645:Underwater camouflage
2640:Structural coloration
2618:Disruptive coloration
2321:Underwater camouflage
2100:Aggressive/Wicklerian
1817:10.1242/dev.116.1.103
1513:Chaetodon capistratus
1177:10.1093/jisesa/iez014
685:
652:Further information:
618:Further information:
582:Further information:
514:cell surface receptor
474:
465:Further information:
449:Chaetodon capistratus
445:Foureye butterflyfish
306:spotted mandarin fish
294:foureye butterflyfish
288:Underwater camouflage
258:
139:butterflies and moths
128:
118:
68:'s display feathers.
24:
3132:Transcription factor
2847:Genetic assimilation
2834:Genetic architecture
2666:Blindness in animals
2598:Counter-illumination
2547:Evolution of the eye
2296:Evolutionary ecology
2281:Deception in animals
2275:Dazzled and Deceived
2235:Animal communication
1523:. July 2009 version.
1220:(15–16): 1047–1054.
596:and lasts until the
506:transcription factor
3228:Morphogenetic field
3145:Influential figures
2142:Emsleyan/Mertensian
1948:2022PNAS..11908661M
1761:1999Sci...283..532K
1687:1996Natur.384..236B
1621:2012NatSR...2E.290O
1574:2016MarBd..46..299B
1562:Marine Biodiversity
1487:on 28 December 2015
1441:Petrie, M. (1992).
1357:2019PLoSO..1426136G
1226:2013JNatH..47.1047S
1114:(1497): 1233–1239.
1009:10.7554/eLife.49544
892:2011Sci...331...73P
841:(1569): 1203–1207.
678:Artificial eyespots
559:Cubitus interruptus
453:disruptive eye mask
2917:Genomic imprinting
2613:Deimatic behaviour
2286:Deimatic behaviour
2010:(1). article 430.
1609:Scientific Reports
1291:"Reptile ID guide"
949:Biological Reviews
738:Behavioral Ecology
718:Patterns in nature
688:
525:regulatory circuit
481:
430:Leptailurus serval
330:Deilephila elpenor
272:sexually selecting
264:
135:
123:
35:
3245:
3244:
3178:Eric F. Wieschaus
3140:
3139:
2958:Pattern formation
2862:Fitness landscape
2724:
2723:
2716:Visual perception
2711:Underwater vision
2676:Feature detection
2671:Eyespot apparatus
2630:Eyespot (mimicry)
2578:Animal coloration
2381:Vision in animals
2348:
2347:
2316:Signalling theory
2291:Mimicry#Evolution
2264:Community ecology
2259:Animal coloration
2105:Ant/Myrmecomorphy
1900:(13): 1159–1166.
1755:(5401): 532–534.
1681:(6606): 236–242.
1629:10.1038/srep00290
1325:978-0-226-51823-7
1061:(1611): 845–851.
943:(November 2005).
784:Ruxton, Graeme D.
708:Animal coloration
516:) Hedgehog (Hh).
487:as a result of a
326:elephant hawkmoth
160:Hipparchia semele
155:deimatic displays
27:gladeye bushbrown
3275:
3188:William McGinnis
3157:Richard Lewontin
3152:C. H. Waddington
3024:
3023:
3001:Neutral networks
2751:
2744:
2737:
2728:
2727:
2696:Palpebral (bone)
2529:Schizochroal eye
2450:
2375:
2368:
2361:
2352:
2351:
2340:Category mimicry
2338:
2337:
2179:
2077:
2070:
2063:
2054:
2053:
2048:
2047:
2037:
2019:
1995:
1986:
1985:
1975:
1926:
1920:
1919:
1909:
1885:
1879:
1878:
1868:
1844:
1831:
1830:
1828:
1800:
1781:
1780:
1740:
1715:
1714:
1695:10.1038/384236a0
1670:
1659:
1658:
1648:
1600:
1594:
1593:
1549:
1543:
1542:
1531:
1525:
1524:
1503:
1497:
1496:
1494:
1492:
1477:
1471:
1470:
1447:Animal Behaviour
1438:
1432:
1431:
1408:Animal Behaviour
1403:
1397:
1396:
1386:
1368:
1351:(12): e0226136.
1336:
1330:
1329:
1309:
1303:
1302:
1300:
1298:
1287:
1281:
1280:
1252:
1246:
1245:
1205:
1199:
1198:
1188:
1156:
1150:
1149:
1139:
1102:Bicyclus anynana
1095:
1089:
1088:
1078:
1046:
1040:
1039:
1029:
1011:
987:
981:
980:
937:
928:
927:
875:
869:
868:
858:
826:
820:
819:
817:
811:. Archived from
810:
792:
776:
770:
769:
744:(5): 1264–1271.
733:
671:Bicyclus anynana
638:Bicyclus anynana
607:Bicyclus anynana
441:
422:
403:
384:
361:
341:
321:
223:Many species of
180:Some species of
175:sexual selection
170:Bicyclus anynana
31:Mycalesis patnia
3283:
3282:
3278:
3277:
3276:
3274:
3273:
3272:
3248:
3247:
3246:
3241:
3232:
3211:
3198:Sean B. Carroll
3136:
3068:
3015:
2979:
2931:
2912:Maternal effect
2895:
2828:
2765:
2755:
2725:
2720:
2649:
2561:
2533:
2451:
2442:
2382:
2379:
2349:
2344:
2325:
2218:
2180:
2171:
2086:
2081:
2051:
1996:
1989:
1927:
1923:
1894:Current Biology
1886:
1882:
1845:
1834:
1801:
1784:
1741:
1718:
1671:
1662:
1601:
1597:
1550:
1546:
1532:
1528:
1504:
1500:
1490:
1488:
1479:
1478:
1474:
1439:
1435:
1404:
1400:
1337:
1333:
1326:
1310:
1306:
1296:
1294:
1289:
1288:
1284:
1253:
1249:
1212:(Salticidae)".
1206:
1202:
1157:
1153:
1096:
1092:
1047:
1043:
988:
984:
941:Stevens, Martin
938:
931:
876:
872:
827:
823:
815:
795:Current Zoology
790:
780:Stevens, Martin
777:
773:
734:
730:
726:
704:
680:
665:
656:
650:
622:
620:Hedgehog (gene)
616:
586:
580:
469:
463:
456:
442:
433:
423:
414:
404:
395:
385:
376:
373:Strymon melinus
369:gray hairstreak
362:
353:
342:
333:
322:
290:
284:
253:
229:Geoffroy's cats
214:
120:Polyphemus moth
113:
108:
19:
12:
11:
5:
3281:
3271:
3270:
3265:
3260:
3258:Animal anatomy
3243:
3242:
3237:
3234:
3233:
3231:
3230:
3225:
3219:
3217:
3213:
3212:
3210:
3209:
3208:
3207:
3195:
3190:
3185:
3180:
3175:
3174:
3173:
3162:François Jacob
3159:
3154:
3148:
3146:
3142:
3141:
3138:
3137:
3135:
3134:
3129:
3124:
3119:
3114:
3109:
3104:
3099:
3098:
3097:
3087:
3082:
3076:
3074:
3070:
3069:
3067:
3066:
3061:
3056:
3051:
3046:
3041:
3036:
3030:
3028:
3021:
3017:
3016:
3014:
3013:
3008:
3003:
2998:
2993:
2987:
2985:
2981:
2980:
2978:
2977:
2972:
2967:
2962:
2961:
2960:
2955:
2945:
2939:
2937:
2933:
2932:
2930:
2929:
2924:
2919:
2914:
2909:
2903:
2901:
2897:
2896:
2894:
2893:
2891:Sequence space
2888:
2883:
2878:
2873:
2868:
2859:
2854:
2849:
2844:
2838:
2836:
2830:
2829:
2827:
2826:
2821:
2820:
2819:
2809:
2804:
2799:
2794:
2789:
2784:
2779:
2773:
2771:
2767:
2766:
2754:
2753:
2746:
2739:
2731:
2722:
2721:
2719:
2718:
2713:
2708:
2703:
2698:
2693:
2688:
2683:
2678:
2673:
2668:
2663:
2657:
2655:
2654:Related topics
2651:
2650:
2648:
2647:
2642:
2637:
2632:
2627:
2626:
2625:
2615:
2610:
2605:
2603:Countershading
2600:
2595:
2590:
2585:
2580:
2575:
2569:
2567:
2563:
2562:
2560:
2559:
2554:
2549:
2543:
2541:
2535:
2534:
2532:
2531:
2526:
2521:
2519:Holochroal eye
2516:
2515:
2514:
2509:
2499:
2498:
2497:
2487:
2482:
2477:
2472:
2467:
2461:
2459:
2453:
2452:
2445:
2443:
2441:
2440:
2439:
2438:
2433:
2428:
2418:
2413:
2408:
2403:
2398:
2392:
2390:
2384:
2383:
2378:
2377:
2370:
2363:
2355:
2346:
2345:
2343:
2342:
2330:
2327:
2326:
2324:
2323:
2318:
2313:
2308:
2303:
2298:
2293:
2288:
2283:
2278:
2271:
2266:
2261:
2256:
2251:
2242:
2237:
2232:
2226:
2224:
2223:Related topics
2220:
2219:
2217:
2216:
2211:
2206:
2201:
2196:
2190:
2188:
2182:
2181:
2174:
2172:
2170:
2169:
2164:
2162:In vertebrates
2159:
2154:
2149:
2144:
2139:
2134:
2129:
2128:
2127:
2117:
2112:
2107:
2102:
2096:
2094:
2088:
2087:
2080:
2079:
2072:
2065:
2057:
2050:
2049:
1987:
1921:
1880:
1832:
1782:
1745:Carroll, S. B.
1716:
1660:
1595:
1568:(1): 299–301.
1544:
1526:
1507:Froese, Rainer
1498:
1472:
1433:
1414:(2): 323–331.
1398:
1331:
1324:
1304:
1282:
1269:10.1086/283868
1263:(5): 770–775.
1247:
1200:
1151:
1090:
1041:
982:
955:(4): 573–588.
929:
870:
821:
818:on 2013-12-12.
771:
727:
725:
722:
721:
720:
715:
713:Nazar (amulet)
710:
703:
700:
679:
676:
664:
661:
649:
646:
615:
612:
579:
576:
538:(Hh) pathway,
462:
459:
458:
457:
455:concealing eye
443:
436:
434:
424:
417:
415:
405:
398:
396:
392:Lacerta agilis
386:
379:
377:
363:
356:
354:
343:
336:
334:
323:
316:
283:
280:
260:Indian peafowl
252:
249:
213:
210:
112:
109:
107:
104:
17:
9:
6:
4:
3:
2:
3280:
3269:
3266:
3264:
3261:
3259:
3256:
3255:
3253:
3240:
3235:
3229:
3226:
3224:
3221:
3220:
3218:
3214:
3206:
3205:
3201:
3200:
3199:
3196:
3194:
3191:
3189:
3186:
3184:
3181:
3179:
3176:
3172:
3169:
3168:
3167:
3166:Jacques Monod
3163:
3160:
3158:
3155:
3153:
3150:
3149:
3147:
3143:
3133:
3130:
3128:
3125:
3123:
3120:
3118:
3115:
3113:
3110:
3108:
3105:
3103:
3100:
3096:
3093:
3092:
3091:
3088:
3086:
3083:
3081:
3080:Homeotic gene
3078:
3077:
3075:
3071:
3065:
3062:
3060:
3057:
3055:
3052:
3050:
3047:
3045:
3042:
3040:
3037:
3035:
3032:
3031:
3029:
3025:
3022:
3018:
3012:
3009:
3007:
3004:
3002:
2999:
2997:
2994:
2992:
2989:
2988:
2986:
2982:
2976:
2973:
2971:
2968:
2966:
2963:
2959:
2956:
2954:
2951:
2950:
2949:
2948:Morphogenesis
2946:
2944:
2941:
2940:
2938:
2934:
2928:
2925:
2923:
2920:
2918:
2915:
2913:
2910:
2908:
2905:
2904:
2902:
2898:
2892:
2889:
2887:
2884:
2882:
2879:
2877:
2874:
2872:
2869:
2867:
2863:
2860:
2858:
2855:
2853:
2850:
2848:
2845:
2843:
2840:
2839:
2837:
2835:
2831:
2825:
2822:
2818:
2815:
2814:
2813:
2810:
2808:
2805:
2803:
2800:
2798:
2795:
2793:
2790:
2788:
2785:
2783:
2782:Reaction norm
2780:
2778:
2775:
2774:
2772:
2768:
2764:
2760:
2752:
2747:
2745:
2740:
2738:
2733:
2732:
2729:
2717:
2714:
2712:
2709:
2707:
2704:
2702:
2699:
2697:
2694:
2692:
2689:
2687:
2684:
2682:
2679:
2677:
2674:
2672:
2669:
2667:
2664:
2662:
2661:Animal senses
2659:
2658:
2656:
2652:
2646:
2643:
2641:
2638:
2636:
2633:
2631:
2628:
2624:
2621:
2620:
2619:
2616:
2614:
2611:
2609:
2606:
2604:
2601:
2599:
2596:
2594:
2593:Chromatophore
2591:
2589:
2586:
2584:
2581:
2579:
2576:
2574:
2571:
2570:
2568:
2564:
2558:
2555:
2553:
2550:
2548:
2545:
2544:
2542:
2540:
2536:
2530:
2527:
2525:
2522:
2520:
2517:
2513:
2510:
2508:
2505:
2504:
2503:
2500:
2496:
2493:
2492:
2491:
2490:Mammalian eye
2488:
2486:
2483:
2481:
2478:
2476:
2473:
2471:
2468:
2466:
2465:Arthropod eye
2463:
2462:
2460:
2458:
2454:
2449:
2437:
2434:
2432:
2429:
2427:
2424:
2423:
2422:
2419:
2417:
2414:
2412:
2409:
2407:
2404:
2402:
2399:
2397:
2394:
2393:
2391:
2389:
2385:
2376:
2371:
2369:
2364:
2362:
2357:
2356:
2353:
2341:
2332:
2331:
2328:
2322:
2319:
2317:
2314:
2312:
2309:
2307:
2304:
2302:
2299:
2297:
2294:
2292:
2289:
2287:
2284:
2282:
2279:
2277:
2276:
2272:
2270:
2267:
2265:
2262:
2260:
2257:
2255:
2252:
2250:
2246:
2243:
2241:
2238:
2236:
2233:
2231:
2228:
2227:
2225:
2221:
2215:
2212:
2210:
2207:
2205:
2202:
2200:
2197:
2195:
2192:
2191:
2189:
2187:
2183:
2178:
2168:
2165:
2163:
2160:
2158:
2155:
2153:
2150:
2148:
2145:
2143:
2140:
2138:
2135:
2133:
2130:
2126:
2123:
2122:
2121:
2118:
2116:
2113:
2111:
2108:
2106:
2103:
2101:
2098:
2097:
2095:
2093:
2089:
2085:
2078:
2073:
2071:
2066:
2064:
2059:
2058:
2055:
2045:
2041:
2036:
2031:
2027:
2023:
2018:
2013:
2009:
2005:
2001:
1994:
1992:
1983:
1979:
1974:
1969:
1965:
1961:
1957:
1953:
1949:
1945:
1941:
1937:
1933:
1925:
1917:
1913:
1908:
1903:
1899:
1895:
1891:
1884:
1876:
1872:
1867:
1862:
1858:
1854:
1850:
1843:
1841:
1839:
1837:
1827:
1822:
1818:
1814:
1810:
1806:
1799:
1797:
1795:
1793:
1791:
1789:
1787:
1778:
1774:
1770:
1766:
1762:
1758:
1754:
1750:
1746:
1739:
1737:
1735:
1733:
1731:
1729:
1727:
1725:
1723:
1721:
1712:
1708:
1704:
1700:
1696:
1692:
1688:
1684:
1680:
1676:
1669:
1667:
1665:
1656:
1652:
1647:
1642:
1638:
1634:
1630:
1626:
1622:
1618:
1614:
1610:
1606:
1599:
1591:
1587:
1583:
1579:
1575:
1571:
1567:
1563:
1559:
1557:
1548:
1540:
1536:
1535:Cott, Hugh B.
1530:
1522:
1521:
1516:
1514:
1508:
1502:
1486:
1482:
1476:
1468:
1464:
1460:
1456:
1452:
1448:
1444:
1437:
1429:
1425:
1421:
1417:
1413:
1409:
1402:
1394:
1390:
1385:
1380:
1376:
1372:
1367:
1362:
1358:
1354:
1350:
1346:
1342:
1335:
1327:
1321:
1317:
1316:
1308:
1292:
1286:
1278:
1274:
1270:
1266:
1262:
1258:
1251:
1243:
1239:
1235:
1231:
1227:
1223:
1219:
1215:
1211:
1204:
1196:
1192:
1187:
1182:
1178:
1174:
1170:
1166:
1162:
1155:
1147:
1143:
1138:
1133:
1129:
1125:
1121:
1117:
1113:
1109:
1105:
1103:
1094:
1086:
1082:
1077:
1072:
1068:
1064:
1060:
1056:
1052:
1045:
1037:
1033:
1028:
1023:
1019:
1015:
1010:
1005:
1001:
997:
993:
986:
978:
974:
970:
966:
962:
958:
954:
950:
946:
942:
936:
934:
925:
921:
917:
913:
909:
905:
901:
897:
893:
889:
885:
881:
874:
866:
862:
857:
852:
848:
844:
840:
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3202:
3095:eyeless gene
2991:Evolvability
2965:Segmentation
2952:
2842:Canalisation
2812:Heterochrony
2802:Heritability
2770:Key concepts
2629:
2524:Parietal eye
2470:Compound eye
2311:Polymorphism
2306:Phagomimicry
2273:
2254:Co-evolution
2146:
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1489:. Retrieved
1485:the original
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1295:. Retrieved
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801:(1): 26–36.
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153:, either in
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93:Antennapedia
70:
62:self-mimicry
51:
42:
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36:
30:
15:
3193:Mike Levine
3102:Distal-less
2927:Polyphenism
2907:Epigenetics
2759:development
2701:Pseudopupil
2583:Aposematism
2502:Mollusc eye
2240:Aposematism
2115:Automimicry
1859:: 253–271.
1811:: 103–109.
1805:Development
1558:(Labridae)"
1453:: 585–586.
584:Distal-less
578:Distal-less
542:(Dll), and
540:Distal-less
388:Sand lizard
365:Automimicry
310:spotted ray
237:pampas cats
233:jungle cats
218:sand lizard
197:automimicry
182:caterpillar
85:Distal-less
41:(sometimes
3252:Categories
3171:Lac operon
2996:Robustness
2975:Modularity
2970:Metamerism
2876:Plasticity
2871:Pleiotropy
2824:Heterotopy
2691:Ommatidium
2623:coincident
2588:Camouflage
2566:Coloration
2507:cephalopod
2401:Chameleons
2245:Camouflage
2214:Vavilovian
2204:Gilbertian
2167:Wasmannian
2092:In animals
1826:1887/11021
724:References
692:Sundarbans
642:B. anynana
628:Drosophila
603:transgenic
572:plasticity
553:Drosophila
531:Drosophila
286:See also:
205:hairstreak
201:Salticidae
193:Lycaenidae
186:Sphingidae
165:camouflage
143:sunbittern
95:, and the
3122:Morphogen
3107:Engrailed
3090:Pax genes
3011:Tinkering
2857:Epistasis
2852:Dominance
2763:phenotype
2706:Rhopalium
2539:Evolution
2512:gastropod
2480:Eye shine
2475:Eagle eye
2406:Dinosaurs
2199:Dodsonian
2186:In plants
2152:MĂĽllerian
2125:Locomotor
2026:2399-3642
1964:0027-8424
1637:2045-2322
1491:23 August
1375:1932-6203
1171:(1): 25.
1128:0962-8452
1018:2050-084X
663:Evolution
544:engrailed
521:evolution
501:diffusion
407:Pygmy owl
346:puss moth
344:Larva of
324:Larva of
298:predators
276:pygmy owl
81:Engrailed
3085:Hox gene
3073:Elements
3054:Homeobox
2573:Albinism
2194:Bakerian
2137:Chemical
2120:Batesian
2044:32770111
1982:35169073
1916:15242612
1875:25341098
1703:12809139
1655:22375251
1537:(1940).
1520:FishBase
1467:53167596
1428:53201236
1393:31851714
1345:PLOS ONE
1297:11 April
1277:34146954
1242:84454608
1195:30794728
1146:12065039
1085:17251116
1036:32041684
977:24868603
969:16221330
924:20443102
916:21212355
908:40986635
865:16024383
786:(2014).
758:19754706
702:See also
614:Hedgehog
536:hedgehog
251:In birds
89:Hedgehog
58:predator
45:) is an
3268:Mimicry
3216:Debates
3027:Systems
2953:Eyespot
2817:Neoteny
2635:Mimicry
2608:Crypsis
2421:Mammals
2249:Crypsis
2147:Eyespot
2084:Mimicry
2035:7414152
1973:8872758
1944:Bibcode
1777:9915699
1757:Bibcode
1749:Science
1711:3341270
1683:Bibcode
1646:3289039
1617:Bibcode
1615:: 290.
1590:8456367
1570:Bibcode
1384:6919575
1353:Bibcode
1222:Bibcode
1186:6390274
1137:1691026
1076:2093980
1027:7012602
888:Bibcode
880:Science
856:1564111
766:1832497
548:Patched
499:source/
496:cautery
282:In fish
268:peacock
262:display
241:servals
147:mimicry
131:Io moth
66:peacock
54:mimicry
43:ocellus
39:eyespot
3117:Ligand
2797:Operon
2426:horses
2388:Vision
2333:
2157:Sexual
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1675:Nature
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239:, and
2495:human
2416:Toads
2396:Birds
2132:Brood
1942:(8).
1707:S2CID
1586:S2CID
1463:S2CID
1424:S2CID
1273:S2CID
1238:S2CID
996:eLife
973:S2CID
920:S2CID
904:JSTOR
816:(PDF)
791:(PDF)
762:S2CID
648:Notch
598:pupal
594:larva
2757:The
2457:Eyes
2436:cats
2431:dogs
2411:Fish
2040:PMID
2022:ISSN
1978:PMID
1960:ISSN
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1633:ISSN
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