241:
183:
480:(the sister group to the remaining Arthropoda) are less well understood. It has been shown that homologs of many eye patterning genes are variably expressed in the eyes of different spider species, but the functional significance of these changes in expression is not well understood, due to lack of functional data. In addition, it has been shown in horseshoe crabs and spiders that Pax6 homologs are not expressed in the same way as their counterparts in insects, suggesting that Pax6 may not be required as a top-level eye patterning switch in chelicerates. Most of the functional data on eye patterning in Chelicerata is drawn from the daddy-longlegs
229:
207:
534:: a stem group lineage that diverged soon before the first true arthropods. The eyes of these creatures are attached to the brain using nerves which enter into the centre of the brain, and there is only one area of the brain devoted to vision. This is similar to the wiring of the median ocelli (small simple eyes) possessed by many arthropods; the eyes also follow a similar pathway through the early development of organisms. This suggests that onychophoran eyes are derived from simple ocelli, and the absence of other eye structures implies that the ancestral arthropod lacked compound eyes, and only used median ocelli to sense light and dark.
43:
2218:
610:
768:
mistaken for nostrils. A further simple eye is located beneath these, on the underside of the carapace; this eye is initially paired during embryonic stages and fuses later in development. A further pair of simple eyes are positioned just in front of the mouth. The simple eyes are probably important during the embryonic or larval stages of the organism, with the compound eyes and median ocelli becoming the dominant sight organs during adulthood. These ocelli are less complex, and probably less derived, than those of the
253:
59:
832:
538:
589:
781:
734:
752:; its eyes are believed to represent the ancestral condition because they have changed so little over evolutionary time. Most other living chelicerates have lost their lateral compound eyes, evolving simple eyes in their place that vary in number. Up to five pairs of lateral eyes occur in scorpions, whereas three pairs of lateral eyes are typical for
240:
772:. Unlike the compound eyes of trilobites, those of horseshoe crabs are triangular in shape; they also have a generative region at their base, but this elongates with time. Hence the one ommatidium at the apex of the triangle was the original "eye" of the larval organism, with subsequent rows added as the organism grew.
566:
There were probably only a single pair of ocelli in the arthropod concestor, since
Cambrian lobopod fossils display a single pair. And while many arthropods today have three, four, or even six, the lack of a common pathway suggests that a pair is the most probable ancestral state. The crustaceans and
554:
A conflicting view notes, however, that compound eyes appeared in many early arthropods, including the trilobites and eurypterids. That suggests that the compound eye may have developed after the onychophoran and arthropod lineages split, but before the radiation of arthropods. This view is supported
815:
eye of most crustaceans, several groups have larvae with simple or compound lateral eyes. The compound eyes of adults develop in a region of the head separate from the region in which the larval median eye develops. New ommatidia are added in semicircular rows at the rear of the eye; during the
362:
spend most of their adult lives on the surface of water, and have their two compound eyes split into four halves, two for underwater vision and two for vision in air. Again, particularly in some
Diptera, ommatidia in different regions of the holoptic male eye may differ visibly in size; the upper
767:
Horseshoe crabs have two large compound eyes on the sides of its head. An additional simple eye is positioned at the rear of each of these structures. In addition to these obvious structures, it also has two smaller ocelli situated in the middle-front of its carapace, which may superficially be
574:
It is deemed probable that the compound eye arose as a result of the 'duplication' of individual ocelli. In turn, the dispersal of compound eyes seems to have created large networks of seemingly independent eyes in some arthropods, such as the larvae of certain insects. In some other insects and
164:
larvae) they can possess resolving powers of 4 degrees of arc, be polarization sensitive and capable of increasing their absolute sensitivity at night by a factor of 1,000 or more. Flying insects can remain level with either type of eye surgically removed, but the two types combine to give
347:, in which seeing prey from particular ommatidia in both compound eyes at the same time, indicates that it is in the right position to snatch in a close-range ambush. Their eyes accordingly are placed in a good position for all-round vision, plus particular concentration on the
658:
was the most common and most primitive. It consisted of many small lenses – between 100 and 15,000 – covered by a single corneal membrane. This was the most ancient kind of eye. This eye morphology was found in the
Cambrian trilobites (the earliest) and survived until the
252:
275:. Examples include most insects, and most of the larger species of Crustacea, such as crabs. Many other organisms, such as vertebrates and Cephalopoda are similarly and analogously dichoptic, which is the common state in animals that are members of the
147:
Most arthropods have at least one of two types of eye: lateral compound eyes, and smaller median ocelli, which are simple eyes. When both are present, the two eye types are used in concert because each has its own advantage. Some insect
319:
and crowding the ocelli, if any. Though technically such eyes still may be regarded dichoptic, the result in the extreme case is that borders of such eyes meet, effectively forming a cap over most of the head. Such an anatomy is called
123:, but the type and origin of this eye varies between groups, and some taxa have secondarily developed simple eyes. The organ's development through the lineage can be estimated by comparing groups that branched early, such as the
529:
No fossil organisms have been identified as similar to the last common ancestor of arthropods; hence the eyes possessed by the first arthropod remains a matter of conjecture. The largest clue into their appearance comes from the
505:
are currently thought to fall within the
Crustacean crown group; while molecular work paved the way for this association, their eye morphology and development is also markedly similar. The eyes are strikingly different from the
816:
first phase of growth, this leads to individual ommatidia being square, but later in development they become hexagonal. The hexagonal pattern will become visible only when the carapace of the stage with square eyes is molted.
206:
228:
182:
279:
and have functionally elaborate eyes. However, there are variations on that scheme. In some groups of animals whose ancestors originally were dichoptic, the eyes of modern species may be crowded together in the
370:
Compound eyes are often not completely symmetrical in terms of ommatidia count. For example, asymmetries have been indicated in honeybees and various flies. This asymmetry has been correlated with behavioural
246:
Stalked dichoptic eyes of a River Crab are typical of mature larger
Crustacea. The reflection of the photographer in different regions of the surface of each eye indicate the basis for stereoscopic vision
823:
occur in some species of crustaceans and some insects, only some of the
Crustacea, such as crabs, bear their eyes on articulated peduncles that permit the eyes to be folded out of the way of trouble.
748:
has traditionally been used in investigations into the eye, because it has relatively large ommatidia with large nerve fibres (making them easy to experiment on). It also falls nears the base of the
339:
In contrast, the need for particular functions may not require extremely large eyes, but do require great resolution and good stereoscopic vision for precise attacks. Good examples may be seen in the
850:
have a compound eye, which is composed of facets and not, as earlier interpretations had it, of clustered stemmata. that were thought to grow in rows, inserted between existing rows of ocelli.
513:
Both ocelli and compound eyes were probably present in the last common arthropod ancestor, and may be apomorphic with ocelli in other phyla, such as the annelids. Median ocelli are present in
801:. This is consistent with the observation that their eyes develop in a very similar fashion. While most crustacean and some insect larvae possess only simple median eyes, such as the
1670:
Gainett, Guilherme; Klementz, Benjamin C.; Blaszczyk, Pola; Setton, Emily V.W.; Murayama, Gabriel P.; Willemart, Rodrigo; Gavish-Regev, Efrat; Sharma, Prashant P. (February 2024).
165:
better performance. Ocelli can detect lower light levels, and have a faster response time, while compound eyes are better at detecting edges and are capable of forming images.
1387:
S. Wehrhahn, William A. Zuker; Harris, WA; Kirschfeld, K; Wehrhahn, C; Zuker, CS (1988), "Targeted misexpression of a
Drosophila opsin gene leads to altered visual function",
367:
the effect is so exaggerated that the upper part of the eye is elevated like a risen cupcake, while its lower part that serves for routine vision looks like a separate organ.
1776:"Ancient mechanisms of visual sense organ development based on comparison of the gene networks controlling larval eye, ocellus, and compound eye specification in Drosophila"
271:
Most species of
Arthropoda with compound eyes bear just two eyes that are located separately and symmetrically, one on each side of the head. This arrangement is called
405:). Together, these genes drive the proliferation of cells early in eye development. Loss of either of these genes results in failure of eye formation. The activity of
355:), showing which ommatidia are covering that field of view; from any position on the median plane, and nowhere else, the two dark spots are symmetrical and identical.
389:(the best-studied arthropod species with respect to developmental biology), among the most important genes for patterning the eyes of insects are the Pax6 homologs
725:. This form of eye consisted of up to 70 much smaller lenses. The cornea separated each lens, and the sclera on each lens terminated on top of each cornea.
710:
eye designs appear haphazard and irregular – possibly constrained by the geometrical complications of packing identical sized lenses on a curved surface. Later
844:
Most myriapods bear stemmata – single lensed eyes which are thought to have evolved by the reduction of a compound eye. However, members of the chilopod genus
358:
Sometimes the needs for visual acuity in different functions conflict, and different parts of the eyes may be adapted to separate functions; for example, the
1437:
Schomburg, Christoph; Turetzek, Natascha; Schacht, Magdalena Ines; Schneider, Julia; Kirfel, Phillipp; Prpic, Nikola-Michael; Posnien, Nico (December 2015).
490:
plays a conserved role in patterning both the visual systems of this species (an example of conservation of gene function, with respect to insects) and that
2142:
649:
eyes. The eye morphology of trilobites is useful for inferring their mode of life, and can function as indicators of the palaeo-environment conditions.
311:, which in turn demands larger compound eyes. The result is that the eyes occupy most of the available surface of the head, reducing the area of the
2326:
1863:
2217:
1949:
901:
are about 5000 times more sensitive than apposition compound eyes. They can, for instance, respond to the position of the full moon.
690:; they were more powerful, with overlapping visual fields, and were particularly useful for nocturnal vision and possibly for colour and
1113:
Seidl, R.; Kaiser, W. (1981). "Visual field size, binocular domain and the ommatidial array of the compound eyes in worker honey bees".
2445:
1922:
1558:
Blackburn, David C.; Conley, Kevin W.; Plachetzki, David C.; Kempler, Karen; Battelle, Barbara-Anne; Brown, Nadean L. (August 2008).
1961:
1932:
429:), which form a protein complex that regulates the transcription of downstream target genes. Thereafter, the two visual systems of
1498:"Differential expression of retinal determination genes in the principal and secondary eyes of Cupiennius salei Keyserling (1877)"
2392:
1012:
Taylor, Charles P. (1981). "Contribution of compound eyes and ocelli to steering of locusts in flight. I. Behavioural analysis".
2135:
2506:
348:
281:
964:
Mayer, G. (2006). "Structure and development of onychophoran eyes: What is the ancestral visual organ in arthropods?".
2254:
869:
169:
351:
median plane. The individual ommatidia are directed in all directions and accordingly, one may see a dark spot (the
2450:
2128:
2090:) (Chilopoda; Notostigmophora): An ultrastructural re-investigation that adds support to the Mandibulata concept".
1883:
Paulus, H.F. (2000), "Phylogeny of the
Myriapoda-Crustacea-Insecta: a new attempt using photoreceptor structure*",
1839:
702:
that separates it from the surrounding lenses. The multiple lenses for the eye were each constructed from a single
1439:"Molecular characterization and embryonic origin of the eyes in the common house spider Parasteatoda tepidariorum"
42:
2321:
740:
have two primary compound eyes and seven secondary simple eyes. Two of the secondary eyes are on the underside.
447:
which demarcates the segments from the top-middle of the head to the more lateral aspects. The ocelli are in an
2511:
1617:"Coming into clear sight at last: Ancestral and derived events during chelicerate visual system development"
879:
385:
58:
1560:"Isolation and expression of Pax6 and atonal homologues in the American horseshoe crab, Limulus polyphemus"
609:
455:
results in loss of the ocelli, but does not affect the compound eyes. Inversely, the transcription factor
2455:
2414:
2409:
2387:
266:
compound eye in the middle of the head, visible here as the dark spot between the bases of the antennae
2435:
2367:
2316:
2001:
1790:
1731:
1396:
1202:
973:
837:
794:
560:
518:
1722:
Harzsch, S.; Hafner, G. (2006), "Evolution of eye development in arthropods: Phylogenetic aspects",
678:
in modern animals, but a somewhat similar eye structure is found in adult male insects in the order
64:
Although arachnids (like this spider) do not have compound eyes, xiphosurans and many other extinct
802:
831:
555:
if a stem-arthropod position is supported for compound-eye bearing
Cambrian organisms such as the
2382:
2107:
2064:
1695:
1671:
1652:
1420:
1193:
1165:
1130:
1095:
820:
786:
660:
1148:
Beersma, D.G.; et al. (1977). "Retinal lattice, visual field and binocularities in flies".
494:
affects the patterning of lateral eyes, but not median eyes (another example of conservation).
2485:
2480:
2440:
2399:
2347:
2056:
2017:
1957:
1928:
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1806:
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1361:
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989:
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537:
482:
316:
312:
1616:
2465:
2298:
2190:
2170:
2099:
2082:
Müller, C.H.G.; Rosenberg, J.; Richter, S.; Meyer-Rochow, V.B. (2003). "The compound eye of
2048:
2009:
1892:
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457:
359:
780:
588:
575:
myriapods, lateral ocelli appear to have arisen by the reduction of lateral compound eyes.
234:
In some male mayflies the eyes are split into separate organs for distinct visual functions
2249:
2185:
2175:
1752:
1672:"Vestigial organs alter fossil placements in an ancient group of terrestrial chelicerates"
293:
214:
190:
31:
1775:
1039:
Meyer-Rochow, V.B. (1974). "Structure and function of the larval eye of the sawfly larva
2036:
2005:
1794:
1735:
1400:
1296:
1206:
977:
937:
288:. In extreme cases such eyes may fuse, effectively into a single eye, as in some of the
2372:
2288:
2276:
2180:
1592:
1559:
1532:
1497:
1473:
1438:
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1247:
1223:
1184:
874:
753:
745:
737:
654:
636:
594:
128:
1897:
1344:
encodes a nuclear protein required for normal eye and leg development in Drosophila".
2500:
2362:
2281:
2259:
2195:
2157:
1918:
1699:
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1056:
548:
444:
435:
304:
285:
2068:
1134:
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2239:
2111:
1424:
1169:
864:
733:
679:
543:
198:
136:
101:
79:
1099:
567:
insects mainly have three ocelli, suggesting that such a formation was present in
2470:
2352:
2271:
2200:
2165:
2052:
859:
769:
749:
599:
531:
514:
477:
352:
344:
329:
153:
124:
109:
105:
65:
49:
1214:
112:
appear to have evolved compound eyes from simple eyes in the opposite fashion.
86:
groups, although they may have evolved more than once within this phylum. Some
2460:
2357:
2205:
2103:
2013:
1802:
1743:
1683:
1514:
1455:
1263:
985:
807:
556:
372:
308:
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2475:
2308:
2264:
2244:
1357:
1319:
846:
798:
761:
722:
721:
is the third eye morphology of trilobites, but it has found only within the
671:
632:
510:, which were traditionally considered to be a sister group to the Hexapoda.
507:
333:
325:
276:
116:
83:
2021:
1810:
1761:
1691:
1648:
1632:
1601:
1541:
1482:
1281:
1232:
993:
52:
have compound eyes, with the exception of some myriapods that have stemmata
1855:
1416:
1373:
1327:
1064:
945:
222:
compound eyes, with the dorsal ommatidia larger than the ventral ommatidia
78:
are the most common form of eye, and are presumably the ancestral form of
17:
2342:
502:
289:
157:
91:
1992:(Xiphosura, Chelicerata) and their afferent and efferent projections.",
694:. Schizochroal eyes have up to 700 large lenses (large compared to
2404:
2377:
1575:
1161:
1126:
1091:
1025:
924:
Land, Michael F.; Fernald, Russell D. (1992), "The Evolution of Eyes",
703:
620:
465:) is required for the patterning of compound eyes, but mutants lacking
259:
96:
87:
1185:"Asymmetric ommatidia count and behavioural lateralization in the ant
1408:
898:
757:
699:
433:
are patterned differently. Anterior head patterning is controlled by
364:
340:
161:
132:
2120:
1340:
Mardon, G.; Solomon, N.M., N. M.; Rubin, G. M., G. M. (Dec 1994). "
1297:"Pattern formation in Drosophila head development: the role of the
1078:
Wilson, M. (1978). "The functional organisation of locust ocelli".
2430:
1496:
Samadi, Leyli; Schmid, Axel; Eriksson, Bo Joakim (December 2015).
779:
732:
470:
149:
262:
and other small Crustacea have their eyes combined into a single
30:"Fly eye" redirects here. For Calvin Harris's record label, see
2124:
2226:
120:
75:
1885:
Journal of Zoological Systematics & Evolutionary Research
698:
lenses). Each lens has a cornea, and each has an individual
674:(Ordovician-Silurian). There is no exact counterpart to the
160:. These eyes usually provide only a rough image, but (as in
413:
includes the activation of the retinal determination genes
100:, the horseshoe crab, and there are suggestions that other
1952:. In Tanacredi, J.T.; Botton, M.L.; Smith, D. (eds.).
303:
On the other hand, some modes of life demand enhanced
94:
also have apposition eyes. They are also possessed by
2035:
Miether, Sebastian T.; Dunlop, Jason A. (July 2016).
1840:"Evolution of eye structure and arthropod phylogeny"
307:, which in compound eyes demands a larger number of
2423:
2335:
2307:
2225:
2156:
521:; lateral ocelli are also present in chelicerates.
670:was found only in one sub-order of trilobite, the
108:by reduction from a compound starting point. Some
797:within the Crustacea, and that the Crustacea are
363:ommatidia tend to be larger. In the case of some
559:. Yet another alternative is that compound eyes
1983:
1981:
1979:
1977:
1975:
1973:
1913:
1911:
1909:
1907:
156:, have a different type of simple eye known as
1717:
1715:
1713:
1711:
1709:
296:. One term for such an arrangement of eyes is
2136:
1988:Battelle, B.A. (December 2006), "The eyes of
1878:
1876:
1874:
1833:
1831:
1829:
1827:
1825:
1823:
1821:
1819:
1252:Current Opinion in Genetics & Development
469:do not exhibit loss of the ocelli. Different
8:
1007:
1005:
1003:
1954:Biology and Conservation of Horseshoe Crabs
959:
957:
955:
2143:
2129:
2121:
919:
917:
324:. Spectacular examples may be seen in the
1896:
1751:
1591:
1531:
1513:
1472:
1454:
1271:
1248:"Pattern formation in the Drosophila eye"
1222:
473:are used in the ocelli of compound eyes.
2037:"Lateral eye evolution in the arachnids"
1924:Invertebrate Palaeontology and Evolution
830:
536:
175:Anatomical distribution of compound eyes
1927:(4th ed.). John Wiley & Sons.
913:
891:
178:
131:to the advanced eye condition found in
2327:Evolution of color vision in primates
1994:Arthropod Structure & Development
1783:Arthropod Structure & Development
1724:Arthropod Structure & Development
1553:
1551:
966:Arthropod Structure & Development
561:independently evolved, multiple times
27:Visual organs possessed by arthropods
7:
686:eyes developed as an improvement on
284:plane; examples include many of the
1615:Friedrich, Markus (December 2022).
938:10.1146/annurev.ne.15.030192.000245
486:, which has been used to show that
1246:Carthew, Richard W (August 2007).
25:
1898:10.1046/j.1439-0469.2000.383152.x
1295:Royet, J; Finkelstein, R (1995),
1080:Journal of Comparative Physiology
188:Many insects, such as the female
2216:
1183:Hunt, E.R.; et al. (2018).
608:
587:
328:and various flies, such as some
251:
239:
227:
205:
181:
57:
41:
1838:Bitsch, C.; Bitsch, J. (2005),
1956:. Springer. pp. 223–235.
1753:11858/00-001M-0000-0012-A87C-4
714:eyes had size graduated lens.
1:
926:Annual Review of Neuroscience
793:It is generally thought that
451:-rich area and disruption of
1057:10.1016/0022-1910(74)90087-0
1045:Journal of Insect Physiology
635:came in three forms, called
2255:Simple eye in invertebrates
2053:10.13156/arac.2006.17.2.103
1950:"Vision in horseshoe crabs"
870:Simple eye in invertebrates
170:Simple eye in invertebrates
2528:
2451:Infrared sensing in snakes
1215:10.1038/s41598-018-23652-4
167:
29:
2322:Evolution of color vision
2214:
2104:10.1007/s00435-003-0085-0
2014:10.1016/j.asd.2006.07.002
1966:– via Google Books.
1937:– via Google Books.
1803:10.1016/j.asd.2006.08.010
1744:10.1016/j.asd.2006.08.009
1684:10.1016/j.cub.2024.02.011
1515:10.1186/s13227-015-0010-x
1456:10.1186/s13227-015-0011-9
1264:10.1016/j.gde.2007.05.001
986:10.1016/j.asd.2006.06.003
819:Although stalked eyes on
82:. They are found in all
375:in ants (turning bias).
1358:10.1242/dev.120.12.3473
1320:10.1242/dev.121.11.3561
880:Optic lobe (arthropods)
776:Insects and crustaceans
563:within the arthropods.
386:Drosophila melanogaster
292:, notably in the genus
1633:10.1002/bies.202200163
1564:Developmental Dynamics
1187:Temnothorax albipennis
841:
790:
741:
551:
476:The visual systems of
119:ancestrally possessed
2456:Monocular deprivation
2415:Underwater camouflage
2410:Structural coloration
2388:Disruptive coloration
2084:Scutigera coleoptrata
1948:Barlow, R.B. (2009).
1856:10.1201/9781420037548
1774:Friedrich, M (2006),
834:
783:
736:
540:
168:Further information:
2507:Arthropod morphology
2436:Blindness in animals
2368:Counter-illumination
2317:Evolution of the eye
838:Lithobius forficatus
544:Anomalocaris daleyae
2006:2006ArtSD..35..261B
1990:Limulus polyphemus
1795:2006ArtSD..35..357F
1736:2006ArtSD..35..319H
1401:1988Natur.333..737F
1207:2018NatSR...8.5825H
978:2006ArtSD..35..231M
795:insects are a clade
194:, shown here, have
2383:Deimatic behaviour
1576:10.1002/dvdy.21634
1194:Scientific Reports
1162:10.1007/bf00656634
1127:10.1007/bf00606065
1092:10.1007/bf00661380
1026:10.1242/jeb.93.1.1
842:
791:
787:Triatoma infestans
742:
661:Permian extinction
552:
541:Fossilised eye of
143:Eyes and functions
2494:
2493:
2486:Visual perception
2481:Underwater vision
2446:Feature detection
2441:Eyespot apparatus
2400:Eyespot (mimicry)
2348:Animal coloration
2151:Vision in animals
1865:978-0-8493-3498-6
1844:Crustacean Issues
1352:(12): 3473–3486.
1314:(11): 3561–3572,
666:The more complex
483:Phalangium opilio
383:In the fruit fly
16:(Redirected from
2519:
2466:Palpebral (bone)
2299:Schizochroal eye
2220:
2145:
2138:
2131:
2122:
2116:
2115:
2089:
2079:
2073:
2072:
2032:
2026:
2025:
1985:
1968:
1967:
1945:
1939:
1938:
1915:
1902:
1901:
1900:
1880:
1869:
1868:
1835:
1814:
1813:
1780:
1771:
1765:
1764:
1755:
1719:
1704:
1703:
1667:
1661:
1660:
1627:(12): e2200163.
1612:
1606:
1605:
1595:
1570:(8): 2209–2219.
1555:
1546:
1545:
1535:
1517:
1493:
1487:
1486:
1476:
1458:
1434:
1428:
1427:
1409:10.1038/333737a0
1395:(6175): 737–41,
1384:
1378:
1377:
1337:
1331:
1330:
1305:
1292:
1286:
1285:
1275:
1243:
1237:
1236:
1226:
1180:
1174:
1173:
1150:J. Comp. Physiol
1145:
1139:
1138:
1115:J. Comp. Physiol
1110:
1104:
1103:
1075:
1069:
1068:
1051:(8): 1565–1591.
1036:
1030:
1029:
1009:
998:
997:
961:
950:
949:
921:
902:
896:
719:abathochroal eye
692:depth perception
676:schizochroal eye
668:schizochroal eye
616:Schizochroal eye
612:
591:
379:Genetic controls
255:
243:
231:
209:
185:
104:developed their
61:
48:Insects and all
45:
21:
2527:
2526:
2522:
2521:
2520:
2518:
2517:
2516:
2512:Vision by taxon
2497:
2496:
2495:
2490:
2419:
2331:
2303:
2221:
2212:
2152:
2149:
2119:
2087:
2081:
2080:
2076:
2034:
2033:
2029:
1987:
1986:
1971:
1964:
1947:
1946:
1942:
1935:
1917:
1916:
1905:
1882:
1881:
1872:
1866:
1837:
1836:
1817:
1778:
1773:
1772:
1768:
1721:
1720:
1707:
1676:Current Biology
1669:
1668:
1664:
1614:
1613:
1609:
1557:
1556:
1549:
1495:
1494:
1490:
1436:
1435:
1431:
1386:
1385:
1381:
1339:
1338:
1334:
1303:
1294:
1293:
1289:
1245:
1244:
1240:
1182:
1181:
1177:
1147:
1146:
1142:
1112:
1111:
1107:
1077:
1076:
1072:
1038:
1037:
1033:
1011:
1010:
1001:
963:
962:
953:
923:
922:
915:
911:
906:
905:
897:
893:
888:
856:
829:
778:
738:Horseshoe crabs
731:
706:crystal. Early
629:
628:
627:
626:
625:
613:
605:
604:
592:
581:
527:
500:
431:D. melanogaster
399:twin of eyeless
381:
267:
256:
247:
244:
235:
232:
223:
215:Tabanus lineola
210:
201:
191:Tabanus lineola
186:
177:
172:
145:
76:Apposition eyes
73:
72:
71:
70:
69:
62:
54:
53:
46:
35:
32:Fly Eye Records
28:
23:
22:
15:
12:
11:
5:
2525:
2523:
2515:
2514:
2509:
2499:
2498:
2492:
2491:
2489:
2488:
2483:
2478:
2473:
2468:
2463:
2458:
2453:
2448:
2443:
2438:
2433:
2427:
2425:
2424:Related topics
2421:
2420:
2418:
2417:
2412:
2407:
2402:
2397:
2396:
2395:
2385:
2380:
2375:
2373:Countershading
2370:
2365:
2360:
2355:
2350:
2345:
2339:
2337:
2333:
2332:
2330:
2329:
2324:
2319:
2313:
2311:
2305:
2304:
2302:
2301:
2296:
2291:
2289:Holochroal eye
2286:
2285:
2284:
2279:
2269:
2268:
2267:
2257:
2252:
2247:
2242:
2237:
2231:
2229:
2223:
2222:
2215:
2213:
2211:
2210:
2209:
2208:
2203:
2198:
2188:
2183:
2178:
2173:
2168:
2162:
2160:
2154:
2153:
2150:
2148:
2147:
2140:
2133:
2125:
2118:
2117:
2098:(4): 191–209.
2088:Linnaeus, 1758
2074:
2047:(2): 103–119.
2027:
1969:
1962:
1940:
1933:
1919:Clarkson, Euan
1903:
1891:(3): 189–208,
1870:
1864:
1815:
1789:(4): 357–378,
1766:
1730:(4): 319–340,
1705:
1662:
1607:
1547:
1488:
1429:
1379:
1332:
1301:homeobox gene"
1287:
1258:(4): 309–313.
1238:
1201:(5825): 5825.
1175:
1156:(3): 207–220.
1140:
1105:
1086:(4): 297–316.
1070:
1031:
999:
972:(4): 231–245.
951:
912:
910:
907:
904:
903:
890:
889:
887:
884:
883:
882:
877:
875:Vision in fish
872:
867:
862:
855:
852:
828:
825:
777:
774:
754:Tetrapulmonata
746:horseshoe crab
730:
729:Horseshoe crab
727:
655:holochroal eye
614:
607:
606:
595:Holochroal eye
593:
586:
585:
584:
583:
582:
580:
579:Trilobite eyes
577:
526:
523:
499:
496:
380:
377:
373:lateralization
269:
268:
257:
250:
248:
245:
238:
236:
233:
226:
224:
211:
204:
202:
187:
180:
176:
173:
144:
141:
129:horseshoe crab
63:
56:
55:
47:
40:
39:
38:
37:
36:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
2524:
2513:
2510:
2508:
2505:
2504:
2502:
2487:
2484:
2482:
2479:
2477:
2474:
2472:
2469:
2467:
2464:
2462:
2459:
2457:
2454:
2452:
2449:
2447:
2444:
2442:
2439:
2437:
2434:
2432:
2431:Animal senses
2429:
2428:
2426:
2422:
2416:
2413:
2411:
2408:
2406:
2403:
2401:
2398:
2394:
2391:
2390:
2389:
2386:
2384:
2381:
2379:
2376:
2374:
2371:
2369:
2366:
2364:
2363:Chromatophore
2361:
2359:
2356:
2354:
2351:
2349:
2346:
2344:
2341:
2340:
2338:
2334:
2328:
2325:
2323:
2320:
2318:
2315:
2314:
2312:
2310:
2306:
2300:
2297:
2295:
2292:
2290:
2287:
2283:
2280:
2278:
2275:
2274:
2273:
2270:
2266:
2263:
2262:
2261:
2260:Mammalian eye
2258:
2256:
2253:
2251:
2248:
2246:
2243:
2241:
2238:
2236:
2235:Arthropod eye
2233:
2232:
2230:
2228:
2224:
2219:
2207:
2204:
2202:
2199:
2197:
2194:
2193:
2192:
2189:
2187:
2184:
2182:
2179:
2177:
2174:
2172:
2169:
2167:
2164:
2163:
2161:
2159:
2155:
2146:
2141:
2139:
2134:
2132:
2127:
2126:
2123:
2113:
2109:
2105:
2101:
2097:
2093:
2092:Zoomorphology
2085:
2078:
2075:
2070:
2066:
2062:
2058:
2054:
2050:
2046:
2042:
2038:
2031:
2028:
2023:
2019:
2015:
2011:
2007:
2003:
2000:(4): 261–74,
1999:
1995:
1991:
1984:
1982:
1980:
1978:
1976:
1974:
1970:
1965:
1963:9780387899589
1959:
1955:
1951:
1944:
1941:
1936:
1934:9781444313321
1930:
1926:
1925:
1920:
1914:
1912:
1910:
1908:
1904:
1899:
1894:
1890:
1886:
1879:
1877:
1875:
1871:
1867:
1861:
1857:
1853:
1849:
1845:
1841:
1834:
1832:
1830:
1828:
1826:
1824:
1822:
1820:
1816:
1812:
1808:
1804:
1800:
1796:
1792:
1788:
1784:
1777:
1770:
1767:
1763:
1759:
1754:
1749:
1745:
1741:
1737:
1733:
1729:
1725:
1718:
1716:
1714:
1712:
1710:
1706:
1701:
1697:
1693:
1689:
1685:
1681:
1677:
1673:
1666:
1663:
1658:
1654:
1650:
1646:
1642:
1638:
1634:
1630:
1626:
1622:
1618:
1611:
1608:
1603:
1599:
1594:
1589:
1585:
1581:
1577:
1573:
1569:
1565:
1561:
1554:
1552:
1548:
1543:
1539:
1534:
1529:
1525:
1521:
1516:
1511:
1507:
1503:
1499:
1492:
1489:
1484:
1480:
1475:
1470:
1466:
1462:
1457:
1452:
1448:
1444:
1440:
1433:
1430:
1426:
1422:
1418:
1414:
1410:
1406:
1402:
1398:
1394:
1390:
1383:
1380:
1375:
1371:
1367:
1363:
1359:
1355:
1351:
1347:
1343:
1336:
1333:
1329:
1325:
1321:
1317:
1313:
1309:
1302:
1300:
1299:orthodenticle
1291:
1288:
1283:
1279:
1274:
1269:
1265:
1261:
1257:
1253:
1249:
1242:
1239:
1234:
1230:
1225:
1220:
1216:
1212:
1208:
1204:
1200:
1196:
1195:
1190:
1188:
1179:
1176:
1171:
1167:
1163:
1159:
1155:
1151:
1144:
1141:
1136:
1132:
1128:
1124:
1120:
1116:
1109:
1106:
1101:
1097:
1093:
1089:
1085:
1081:
1074:
1071:
1066:
1062:
1058:
1054:
1050:
1046:
1042:
1035:
1032:
1027:
1023:
1019:
1015:
1008:
1006:
1004:
1000:
995:
991:
987:
983:
979:
975:
971:
967:
960:
958:
956:
952:
947:
943:
939:
935:
931:
927:
920:
918:
914:
908:
900:
895:
892:
885:
881:
878:
876:
873:
871:
868:
866:
863:
861:
858:
857:
853:
851:
849:
848:
840:
839:
833:
826:
824:
822:
817:
814:
810:
809:
804:
803:Bolwig organs
800:
796:
789:
788:
782:
775:
773:
771:
765:
763:
759:
755:
751:
747:
739:
735:
728:
726:
724:
720:
715:
713:
709:
705:
701:
697:
693:
689:
685:
681:
677:
673:
669:
664:
662:
657:
656:
650:
648:
644:
640:
639:
634:
623:
622:
617:
611:
602:
601:
596:
590:
578:
576:
572:
570:
564:
562:
558:
550:
549:Emu Bay Shale
546:
545:
539:
535:
533:
532:onychophorans
524:
522:
520:
516:
511:
509:
504:
497:
495:
493:
489:
485:
484:
479:
474:
472:
468:
464:
460:
459:
454:
450:
446:
445:homeobox gene
442:
438:
437:
436:orthodenticle
432:
428:
424:
420:
416:
412:
408:
404:
400:
396:
392:
388:
387:
378:
376:
374:
368:
366:
365:Ephemeroptera
361:
356:
354:
350:
346:
342:
337:
335:
331:
327:
323:
318:
314:
310:
306:
305:visual acuity
301:
299:
295:
291:
287:
286:Archaeognatha
283:
278:
274:
265:
261:
254:
249:
242:
237:
230:
225:
221:
217:
216:
208:
203:
200:
199:compound eyes
197:
193:
192:
184:
179:
174:
171:
166:
163:
159:
155:
151:
142:
140:
138:
134:
130:
126:
122:
121:compound eyes
118:
113:
111:
107:
103:
99:
98:
93:
89:
85:
81:
77:
67:
60:
51:
44:
33:
19:
2294:Parietal eye
2240:Compound eye
2234:
2095:
2091:
2083:
2077:
2044:
2040:
2030:
1997:
1993:
1989:
1953:
1943:
1923:
1888:
1884:
1847:
1843:
1786:
1782:
1769:
1727:
1723:
1675:
1665:
1624:
1620:
1610:
1567:
1563:
1505:
1501:
1491:
1446:
1442:
1432:
1392:
1388:
1382:
1349:
1345:
1341:
1335:
1311:
1307:
1298:
1290:
1255:
1251:
1241:
1198:
1192:
1186:
1178:
1153:
1149:
1143:
1118:
1114:
1108:
1083:
1079:
1073:
1048:
1044:
1040:
1034:
1017:
1014:J. Exp. Biol
1013:
969:
965:
929:
925:
894:
865:Parietal eye
845:
843:
836:
818:
806:
799:monophyletic
792:
785:
766:
750:chelicerates
743:
718:
716:
712:schizochroal
711:
708:schizochroal
707:
695:
687:
684:Schizochroal
683:
680:Strepsiptera
675:
667:
665:
653:
651:
647:abathochroal
646:
643:schizochroal
642:
637:
631:The eyes of
630:
619:
598:
573:
568:
565:
557:Radiodontids
553:
542:
528:
519:mandibulates
515:chelicerates
512:
501:
491:
487:
481:
475:
466:
462:
456:
452:
448:
440:
434:
430:
426:
422:
418:
414:
410:
406:
402:
398:
394:
390:
384:
382:
369:
357:
338:
321:
302:
297:
272:
270:
263:
219:
213:
195:
189:
154:caterpillars
146:
139:arthropods.
114:
110:caterpillars
102:chelicerates
95:
80:compound eye
74:
66:chelicerates
50:mandibulates
2471:Pseudopupil
2353:Aposematism
2272:Mollusc eye
2041:Arachnology
1850:: 185–214,
1346:Development
1308:Development
932:(1): 1–29,
860:Mollusc eye
770:Mandibulata
600:Paralejurus
571:concestor.
488:eyes absent
478:Chelicerata
423:eyes absent
415:sine oculis
353:pseudopupil
345:Mantispidae
330:Acroceridae
125:velvet worm
106:simple eyes
2501:Categories
2461:Ommatidium
2393:coincident
2358:Camouflage
2336:Coloration
2277:cephalopod
2171:Chameleons
909:References
808:Drosophila
696:holochroal
688:holochroal
638:holochroal
633:trilobites
326:Anisoptera
135:and other
117:arthropods
18:Insect eye
2476:Rhopalium
2309:Evolution
2282:gastropod
2250:Eye shine
2245:Eagle eye
2176:Dinosaurs
2061:2050-9928
1700:267821504
1657:253246397
1641:0265-9247
1621:BioEssays
1584:1058-8388
1524:2041-9139
1508:(1): 16.
1465:2041-9139
1449:(1): 15.
1366:0950-1991
1342:dachshund
1121:: 17–26.
886:Footnotes
847:Scutigera
827:Myriapods
821:peduncles
762:Amblypygi
723:Eodiscina
672:Phacopina
547:from the
508:myriapods
498:Evolution
492:dachshund
458:dachshund
360:Gyrinidae
334:Tabanidae
309:ommatidia
298:cycloptic
277:Bilateria
273:dichoptic
264:cycloptic
212:The male
196:dichoptic
84:arthropod
2343:Albinism
2069:89428386
2022:18089075
1921:(2009).
1811:18089081
1762:18089079
1692:38401545
1649:36317531
1602:18651657
1542:26034575
1483:26034574
1282:17618111
1233:29643429
1135:41150125
1020:: 1–18.
994:18089073
854:See also
813:naupliar
811:and the
503:Hexapods
349:anterior
341:Mantodea
322:holoptic
315:and the
290:Copepoda
260:copepods
220:holoptic
158:stemmata
152:, e.g.,
92:bivalves
88:annelids
2405:Mimicry
2378:Crypsis
2191:Mammals
2112:6466405
2002:Bibcode
1791:Bibcode
1732:Bibcode
1593:2577597
1533:4450993
1502:EvoDevo
1474:4450840
1443:EvoDevo
1425:4248264
1417:2455230
1397:Bibcode
1374:7821215
1328:8582270
1273:2693403
1224:5895843
1203:Bibcode
1170:6384399
1065:4854430
974:Bibcode
946:1575438
835:Eye of
784:Eye of
758:spiders
756:(e.g.,
704:calcite
621:Phacops
391:eyeless
294:Cyclops
137:derived
133:insects
97:Limulus
2196:horses
2158:Vision
2110:
2067:
2059:
2020:
1960:
1931:
1862:
1809:
1760:
1698:
1690:
1655:
1647:
1639:
1600:
1590:
1582:
1540:
1530:
1522:
1481:
1471:
1463:
1423:
1415:
1389:Nature
1372:
1364:
1326:
1280:
1270:
1231:
1221:
1168:
1133:
1100:572458
1098:
1063:
992:
944:
899:Ocelli
700:sclera
645:, and
525:Origin
471:opsins
421:) and
317:vertex
282:median
162:sawfly
150:larvae
2265:human
2186:Toads
2166:Birds
2108:S2CID
2065:S2CID
1779:(PDF)
1696:S2CID
1653:S2CID
1421:S2CID
1304:(PDF)
1166:S2CID
1131:S2CID
1096:S2CID
1041:Perga
569:their
443:), a
313:frons
258:Many
2227:Eyes
2206:cats
2201:dogs
2181:Fish
2057:ISSN
2018:PMID
1958:ISBN
1929:ISBN
1860:ISBN
1807:PMID
1758:PMID
1688:PMID
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