485:
female's tergites have only a narrow pigment stripe. This sexually dimorphic pigmentation pattern is controlled by a genetic regulatory circuit involving the Hox gene Abd-B. Loss-of-function mutations of Abd-B cause the loss of male-specific pigmentation, while gain-of-function alleles, such as Abd-BMcp, cause the expansion of pigmentation to the A4 segment or even to the
25:
492:
Pigmentation of the posterior male abdomen is a trait found in many members of the melanogaster species group but not in several other major groups. The dimorphic regulation of bab expression is closely correlated with dimorphic pigmentation as well other pigmentation patterns. It is not known,
484:
Hox genes have been implicated in the evolution of many animal body patterns. Hox protein directly activates expression of the yellow pigmentation gene in posterior segments. In D. melanogaster, the male has fully pigmented tergites in the fifth and sixth abdominal segments (A5 and A6), whereas the
357:
with a higher efficiency than
Dopamine-chrome. Some authors have proposed that the black pigment in abdominal cuticle was Dopa-melanin produced from Dopa. Incubation of abdominal cuticles or wings of unpigmented pharates with Dopamine is sufficient to produce black pigment, which suggests that this
316:
over-expression induces dark pigmentation in the anterior region of the tergites. However, careful examination revealed that this ectopic pigmentation was not as dark as the normal pigmentation in the posterior region of the tergites. This was more visible in A4 and A5 segments. By contrast, when
374:
females is darker when they develop at low temperature. This is particularly pronounced in posterior abdominal segment. Plasticity of abdominal pigmentation is likely to have functional consequences as abdominal pigmentation has been linked to thermoregulation and resistance to UV,
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In stage A pharates, two cells at the base of bristles expressed y. This expression had a similar intensity when pharates were raised at 18 °C and at 29 °C. These two cells are likely to be the socket and the shaft, the only pigmented cells of the bristle organ. In addition,
440:
expression (A, B and C) based on the degree of maturation of abdominal bristles . These stages correspond approximately to a transition from stage P11(i) to stage P12(ii) as described by
Bainbridge and Bownes with morphological markers at 25 °C.
431:
expression is modulated by temperature in the epidermis of abdominal segments A5, A6 and A7 in female pharates (1.97 fold more expressed at 18 °C than at 29 °C). In order to analyse the spatial expression of y, many researchers performed
475:
was no longer expressed at the base of bristles and the bristles were almost fully pigmented. Furthermore, its overall expression in tergites was reduced compared to stage B and more similar between pharates grown at 18 °C and 29 °C.
402:
abdominal pigmentation are relatively well known, in particular those encoding the enzymes required for the synthesis of cuticle pigments. It has been reported recently that the thermal plasticity of female abdominal pigmentation in
183:
and sclerotin. By mapping the genetic basis of natural variation in body pigmentation, new genes affecting pigment biosynthesis as well as regulatory regions that determine when and where pigmentation will develop were discovered.
178:
pigmentation do not form part of this pathway or any parallel pathway. Furthermore, the genes that lead to natural variation in body pigmentation are not necessarily the same genes that are directly involved in the biosynthesis of
1518:
Walter MF, Zeineh LL, Black BC, McIvor WE, Wright TR, Biessmann H (1996). "Catecholamine metabolism and in vitro induction of premature cuticle melanization in wild type and pigmentation mutants of
Drosophila melanogaster".
1413:
Wittkopp PJ, Stewart EE, Arnold LL, Neidert AH, Haerum BK, Thompson EM, et al. (October 2009). "Intraspecific polymorphism to interspecific divergence: genetics of pigmentation in
Drosophila".
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is sufficient to explain the black pigmentation observed at 18 °C, the researcher increased their expression in abdominal epidermis at 29 °C to mimic the effect of lower temperature.
1318:
Gompel N, Prud'homme B, Wittkopp PJ, Kassner VA, Carroll SB (February 2005). "Chance caught on the wing: cis-regulatory evolution and the origin of pigment patterns in
Drosophila".
449:
was expressed in the posterior region of each tergite in segments A2 to A6. This expression was much broader and stronger in pharates grown at 18 °C compared to 29 °C. In A6,
43:
358:
black pigment is produced from
Dopamine and is therefore Dopamine-melanin. It is also known that Ddc down-regulation leads to a complete loss of black and brown pigments.
1556:"Direct and correlated responses to laboratory selection for body melanisation in Drosophila melanogaster: support for the melanisation-desiccation resistance hypothesis"
565:
Wittkopp PJ, Beldade P (February 2009). "Development and evolution of insect pigmentation: genetic mechanisms and the potential consequences of pleiotropy".
1371:"The expansion of body coloration involves coordinated evolution in cis and trans within the pigmentation regulatory network of Drosophila prostipennis"
2011:
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was expressed in the whole tergite at 18 °C, and only in the posterior region of the tergite at 29 °C. In A7, at 18 °C, the whole tergite expressed
174:
The melanin/sclerotin biosynthetic pathway and its underlying genetic basis have been well studied. However, many of the genes known to affect
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1230:
1954:
Gompel N, Carroll SB (August 2003). "Genetic mechanisms and constraints governing the evolution of correlated traits in drosophilid flies".
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Kopp A, Duncan I, Godt D, Carroll SB (November 2000). "Genetic control and evolution of sexually dimorphic characters in
Drosophila".
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is sex-specifically regulated in the posterior abdomen. Furthermore, the evolution of wing or abdominal pigmentation patterns between
1011:
Wittkopp PJ, Carroll SB, Kopp A (September 2003). "Evolution in black and white: genetic control of pigment patterns in
Drosophila".
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61:
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combined over-expression at 29 °C is necessary and sufficient to reproduce the pigmentation phenotype observed at low temperature.
521:
1213:
Wright TR (1987). "The
Genetics of Biogenic Amine Metabolism, Sclerotization, and Melanization in Drosophila Melanogaster".
1867:"Phenotypic Plasticity through Transcriptional Regulation of the Evolutionary Hotspot Gene tan in Drosophila melanogaster"
239:
is known to be required but not sufficient for black melanin production. Studies have indicated that the black melanin is
251:
at 29 °C is necessary and sufficient to reproduce the black phenotype observed at 18°C. Thus, the stronger expression of
493:
however, which regulatory interactions among Abd-B, bab, dsx, and pigmentation genes are direct and which are indirect.
1814:
Massey JH, Wittkopp PJ (2016). "The
Genetic Basis of Pigmentation Differences within and Between Drosophila Species".
104:
shows enormous phenotypic variation between species, populations, and individuals, and even within individuals during
489:. The sexually dimorphic pigment pattern depends upon regulatory interactions among the Abd-B, bab, and dsx genes.
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is seven times more expressed at 18 °C than at 29 °C in the posterior abdominal epidermis of young adult females.
124:. Changes in pigmentation are often adaptive and vital to the fitness of the organism. Much is known about the
808:"Unraveling the thread of nature's tapestry: the genetics of diversity and convergence in animal pigmentation"
1466:"The evolution of gene regulation underlies a morphological difference between two Drosophila sister species"
117:
90:
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expression was reduced in the socket and the shaft, while the bristle began to be pigmented. Furthermore,
1654:"An Experimental Evolution Test of the Relationship between Melanism and Desiccation Survival in Insects"
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Abdominal pigmentation in Drosophilids represents an appropriate model to dissect the molecular bases of
1143:
Moussian B (May 2010). "Recent advances in understanding mechanisms of insect cuticle differentiation".
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367:
256:
953:"The regulation and evolution of a genetic switch controlling sexually dimorphic traits in Drosophila"
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up-regulation to induce a fully black pigmentation. In order to test whether the strong expression of
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but it is also a relatively simple and easily measured phenotype to study the genetic architecture of
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912:
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Lande R (March 1980). "Sexual Dimorphism, Sexual Selection, and Adaptation in Polygenic Characters".
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353:. Yellow is related to two other enzymes, Yellow-f and Yellow-f2, which can be used as substrate for
94:. It has been used as a model for understanding the development and evolution of morphological
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394:. Furthermore, as abdominal pigmentation is highly evolvable, it has been investigated to study the
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661:"The functional basis of wing patterning in Heliconius butterflies: the molecules behind mimicry"
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Lindgren J, Moyer A, Schweitzer MH, Sjövall P, Uvdal P, Nilsson DE, et al. (August 2015).
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was still more expressed in the abdominal epidermis of pharates grown at 18 °C than at 29 °C.
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1936:
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1048:"Genetics of a difference in pigmentation between Drosophila yakuba and Drosophila santomea"
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Kronforst MR, Barsh GS, Kopp A, Mallet J, Monteiro A, Mullen SP, et al. (July 2012).
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involves transcriptional modulation of the pigmentation gene tan (t). This gene encodes a
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that often has adaptive significance. Pigmentation has extensively been studied in
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1113:
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Kopp A (June 2006). "Basal relationships in the Drosophila melanogaster species group".
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were over-expressed in the dorsal region of the abdomen, the anterior region of the
116:. It also varies between species, contributing to species recognition, mate choice,
1991:
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1605:"Thermoregulatory strategy may shape immune investment in Drosophila melanogaster"
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of female pharates grown at 18 °C or 29 °C and could distinguish three stages of
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generally has a stripe of dark coloration (melanin) on a lighter tan background (
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Williams TM, Selegue JE, Werner T, Gompel N, Kopp A, Carroll SB (August 2008).
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as it is sensitive to temperature in many species. Abdominal pigmentation of
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710:"Gene regulation networks generate diverse pigmentation patterns in plants"
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383:. Abdominal pigmentation is also associated to resistance to desiccation.
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gene is required for the production of black melanin and in the absence of
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and the genes that control the temporal and spatial distribution of this
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is required but not sufficient for production of black pigment. Indeed,
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spatial expression. Temperature also controls the spatial expression of
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10.1002/(sici)1520-6327(1996)31:2<219::aid-arch9>3.0.co;2-u
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bases of morphological variation within species. The genes involved in
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Ordway AJ, Hancuch KN, Johnson W, Wiliams TM, Rebeiz M (August 2014).
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was as black as the posterior border of the tergites. This shows that
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Rajpurohit S, Peterson LM, Orr AJ, Marlon AJ, Gibbs AG (2016-09-22).
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species and has been used as a model to dissect the genetic bases of
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expression associated with bristles is not modulated by temperature.
1263:"Evolution of yellow gene regulation and pigmentation in Drosophila"
860:
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Abdominal pigmentation differs between males and females in several
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Gibert JM, Mouchel-Vielh E, De Castro S, Peronnet F (August 2016).
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of wild-type females and females over-expressing either
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Not only is body pigmentation ecologically relevant in
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that regulate the biochemical synthesis of pigments in
39:
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at a high level, whereas it was much weaker at 29 °C.
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Wittkopp PJ, Vaccaro K, Carroll SB (September 2002).
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Evolution; International Journal of Organic Evolution
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over-expression does not change pigmentation whereas
163:, the epidermal cells underlying the cuticle secrete
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Ramniwas S, Kajla B, Dev K, Parkash R (April 2013).
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may be too technical for most readers to understand
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708:Albert NW, Davies KM, Schwinn KE (2014-06-13).
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349:Yellow gene is required for the production of
167:-derived catecholamines into the cuticle for
120:, protection (warning signals), mimicry, and
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1765:Riedel F, Vorkel D, Eaton S (January 2011).
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567:Seminars in Cell & Developmental Biology
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411:implicated in the production of melanin.
216:species correlates with modifications of
62:Learn how and when to remove this message
46:, without removing the technical details.
308:(UAS-t/+; UAS-y/pnr-Gal4) at 29 °C. The
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341:Production of dopamine-melanin by the
269:gain- of-function must be combined to
108:. It gives rise to natural variation,
905:Molecular Phylogenetics and Evolution
44:make it understandable to non-experts
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812:Pigment Cell & Melanoma Research
1609:The Journal of Experimental Biology
1560:The Journal of Experimental Biology
1065:10.1111/j.0014-3820.2002.tb00150.x
784:10.1146/annurev-ento-010814-020942
259:of female abdominal pigmentation.
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1215:Molecular Genetics of Development
1188:Trends in Ecology & Evolution
659:Kronforst MR, Papa R (May 2015).
526:Journal of Visualized Experiments
2012:Drosophila melanogaster genetics
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616:Proceedings. Biological Sciences
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714:Plant Signaling & Behavior
300:(UAS-t/+; pnr-Gal4/+) or both
224:in the abdominal epidermis of
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1223:10.1016/S0065-2660(08)60008-5
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1933:10.1016/0020-1790(88)90023-6
1884:10.1371/journal.pgen.1006218
1679:10.1371/journal.pone.0163414
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255:at 18°C also contributes to
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1388:10.1016/j.ydbio.2014.05.023
925:10.1016/j.ympev.2006.01.029
771:Annual Review of Entomology
765:Monteiro A (January 2015).
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84:simple but highly variable
16:Genetic trait in Drosophila
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1483:10.1016/j.cell.2008.01.014
1200:10.1016/j.tree.2003.09.006
1165:10.1016/j.ibmb.2010.03.003
1122:10.1016/j.ibmb.2009.10.007
969:10.1016/j.cell.2008.06.052
480:Regulation of pigmentation
75:Abdominal pigmentation in
147:in complex traits. Each
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405:Drosophila melanogaster
372:Drosophila melanogaster
206:Drosophila melanogaster
159:). During pre-and post-
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77:Drosophila melanogaster
628:10.1098/rspb.2015.0614
419:Temperature modulation
1375:Developmental Biology
471:In stage C pharates,
460:In stage B pharates,
434:in-situ hybridization
368:phenotypic plasticity
362:Effect of temperature
502:ectopic pigmentation
1976:10.1038/nature01787
1968:2003Natur.424..931G
1921:Insect Biochemistry
1816:Genes and Evolution
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1670:2016PLoSO..1163414R
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1340:10.1038/nature03235
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1279:2002CBio...12.1547W
1157:2010IBMB...40..363M
1114:2010IBMB...40..166A
917:2006MolPE..39..787K
726:2014PlSiB...9E9526A
273:down-regulation or
171:and melanisation.
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1573:10.1242/jeb.076166
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622:(1813): 20150614.
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257:thermal plasticity
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1777:(1): 149–58.
1776:
1772:
1768:
1761:
1758:
1753:
1749:
1745:
1741:
1737:
1733:
1729:
1725:
1721:
1717:
1710:
1708:
1704:
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1690:
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1659:
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1574:
1569:
1565:
1561:
1557:
1550:
1547:
1542:
1538:
1534:
1530:
1527:(2): 219–33.
1526:
1522:
1514:
1512:
1510:
1506:
1501:
1497:
1493:
1489:
1484:
1479:
1476:(5): 783–93.
1475:
1471:
1467:
1460:
1457:
1452:
1448:
1444:
1440:
1436:
1432:
1428:
1424:
1420:
1416:
1409:
1407:
1403:
1398:
1394:
1389:
1384:
1381:(2): 431–40.
1380:
1376:
1372:
1365:
1362:
1357:
1353:
1349:
1345:
1341:
1337:
1333:
1329:
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1234:
1228:
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1201:
1197:
1193:
1189:
1182:
1179:
1174:
1170:
1166:
1162:
1158:
1154:
1151:(5): 363–75.
1150:
1146:
1139:
1136:
1131:
1127:
1123:
1119:
1115:
1111:
1108:(3): 166–78.
1107:
1103:
1096:
1094:
1092:
1088:
1083:
1079:
1075:
1071:
1066:
1061:
1057:
1053:
1049:
1042:
1039:
1034:
1030:
1026:
1022:
1018:
1014:
1007:
1005:
1003:
1001:
999:
997:
993:
988:
984:
979:
974:
970:
966:
963:(4): 610–23.
962:
958:
954:
947:
945:
943:
939:
934:
930:
926:
922:
918:
914:
911:(3): 787–98.
910:
906:
899:
897:
895:
891:
886:
882:
878:
874:
870:
866:
862:
858:
854:
850:
843:
840:
835:
831:
826:
821:
818:(4): 411–33.
817:
813:
809:
802:
799:
794:
790:
785:
780:
777:(1): 253–71.
776:
772:
768:
761:
758:
753:
749:
744:
739:
735:
731:
727:
723:
720:(9): e29526.
719:
715:
711:
704:
701:
696:
692:
687:
682:
678:
674:
670:
666:
662:
655:
652:
647:
643:
638:
633:
629:
625:
621:
617:
613:
606:
603:
598:
594:
589:
584:
580:
576:
572:
568:
561:
558:
553:
549:
544:
539:
535:
534:10.3791/55732
531:
527:
523:
516:
513:
506:
504:
503:
496:
494:
490:
488:
479:
477:
474:
469:
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463:
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344:
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286:
284:
280:
276:
272:
268:
264:
260:
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254:
250:
246:
242:
238:
234:
231:By contrast,
229:
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199:
191:
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177:
172:
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111:
107:
103:
99:
97:
93:
92:
87:
83:
79:
78:
66:
63:
55:
45:
41:
35:
32:This article
30:
21:
20:
1959:
1955:
1949:
1924:
1920:
1874:
1870:
1860:
1815:
1809:
1774:
1770:
1760:
1719:
1715:
1661:
1657:
1647:
1612:
1608:
1598:
1563:
1559:
1549:
1524:
1520:
1473:
1469:
1459:
1418:
1414:
1378:
1374:
1364:
1323:
1319:
1313:
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1266:
1214:
1208:
1191:
1187:
1181:
1148:
1144:
1138:
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1101:
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908:
904:
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848:
842:
815:
811:
801:
774:
770:
760:
717:
713:
703:
668:
664:
654:
619:
615:
605:
573:(1): 65–71.
570:
566:
560:
525:
515:
500:
491:
483:
472:
470:
465:
461:
459:
454:
450:
446:
443:
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428:
422:
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404:
399:
387:
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371:
365:
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221:
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205:
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197:
195:
189:
175:
173:
152:
140:
138:
134:biosynthesis
129:
102:Pigmentation
100:
89:
76:
74:
73:
58:
49:
33:
1771:Development
671:(1): 1–19.
588:10400.7/197
110:polyphenism
507:References
400:Drosophila
388:Drosophila
355:Dopachrome
228:females.
214:Drosophila
151:of female
141:Drosophila
96:phenotypes
1941:0020-1790
1082:221733289
869:0014-3820
849:Evolution
409:hydrolase
396:molecular
381:parasites
377:pathogens
157:sclerotin
2006:Category
1984:12931186
1903:27508387
1852:27282023
1801:29017593
1793:21138977
1744:11117736
1698:27658246
1658:PLOS ONE
1631:25147243
1590:22483971
1582:23239892
1500:15447569
1492:18329365
1443:19900891
1397:24907418
1356:16422483
1348:15690032
1297:12372246
1173:20347980
1130:19932179
1074:12487356
1033:12957543
987:18724934
933:16527496
885:28563426
834:22578174
793:25341098
752:25763693
695:25953905
665:Genetics
646:26290071
597:18977308
552:28605370
327:tergites
290:cuticles
241:Dopamine
165:tyrosine
106:ontogeny
1992:4415001
1964:Bibcode
1894:4980059
1843:5002358
1724:Bibcode
1689:5033579
1666:Bibcode
1639:6798309
1541:8580497
1451:6796236
1423:Bibcode
1415:Science
1328:Bibcode
1305:2301246
1275:Bibcode
1241:3124532
1153:Bibcode
1110:Bibcode
978:2597198
913:Bibcode
877:2407393
743:4205132
722:Bibcode
686:4423356
637:4632609
543:5608185
425:RT-qPCR
226:pharate
181:melanin
161:ecdysis
149:tergite
122:crypsis
38:Please
1990:
1982:
1956:Nature
1939:
1901:
1891:
1850:
1840:
1830:
1799:
1791:
1752:261526
1750:
1742:
1716:Nature
1696:
1686:
1637:
1629:
1588:
1580:
1539:
1498:
1490:
1449:
1441:
1395:
1354:
1346:
1320:Nature
1303:
1295:
1239:
1229:
1171:
1128:
1080:
1072:
1031:
985:
975:
931:
883:
875:
867:
832:
791:
750:
740:
693:
683:
644:
634:
595:
550:
540:
487:thorax
473:yellow
466:yellow
462:yellow
455:yellow
451:yellow
447:yellow
438:yellow
429:yellow
343:yellow
331:yellow
319:yellow
310:yellow
302:yellow
294:yellow
279:yellow
267:yellow
263:yellow
253:yellow
245:yellow
237:yellow
233:yellow
222:yellow
218:yellow
210:yellow
202:yellow
198:yellow
190:yellow
1988:S2CID
1797:S2CID
1748:S2CID
1635:S2CID
1586:S2CID
1496:S2CID
1447:S2CID
1352:S2CID
1301:S2CID
1078:S2CID
873:JSTOR
427:that
317:both
271:ebony
126:genes
86:trait
80:is a
1980:PMID
1937:ISSN
1899:PMID
1848:PMID
1828:ISBN
1789:PMID
1740:PMID
1694:PMID
1627:PMID
1578:PMID
1537:PMID
1488:PMID
1470:Cell
1439:PMID
1393:PMID
1344:PMID
1293:PMID
1237:PMID
1227:ISBN
1169:PMID
1126:PMID
1070:PMID
1029:PMID
983:PMID
957:Cell
929:PMID
881:PMID
865:ISSN
830:PMID
789:PMID
748:PMID
691:PMID
642:PMID
593:PMID
548:PMID
345:gene
333:and
321:and
304:and
281:and
247:and
196:The
192:gene
188:The
112:and
1972:doi
1960:424
1929:doi
1889:PMC
1879:doi
1838:PMC
1820:doi
1779:doi
1775:138
1732:doi
1720:408
1684:PMC
1674:doi
1617:doi
1613:217
1568:doi
1564:216
1529:doi
1478:doi
1474:132
1431:doi
1419:326
1383:doi
1379:392
1336:doi
1324:433
1283:doi
1219:doi
1196:doi
1161:doi
1118:doi
1060:doi
1021:doi
973:PMC
965:doi
961:134
921:doi
857:doi
820:doi
779:doi
738:PMC
730:doi
681:PMC
673:doi
669:200
632:PMC
624:doi
620:282
583:hdl
575:doi
538:PMC
530:doi
413:tan
379:or
335:tan
323:tan
314:tan
306:tan
298:tan
283:tan
275:tan
249:tan
98:.
42:to
2008::
1986:.
1978:.
1970:.
1958:.
1935:.
1925:18
1923:.
1911:^
1897:.
1887:.
1875:12
1873:.
1869:.
1846:.
1836:.
1826:.
1795:.
1787:.
1773:.
1769:.
1746:.
1738:.
1730:.
1718:.
1706:^
1692:.
1682:.
1672:.
1662:11
1660:.
1656:.
1633:.
1625:.
1611:.
1607:.
1584:.
1576:.
1562:.
1558:.
1535:.
1525:31
1523:.
1508:^
1494:.
1486:.
1472:.
1468:.
1445:.
1437:.
1429:.
1417:.
1405:^
1391:.
1377:.
1373:.
1350:.
1342:.
1334:.
1322:.
1299:.
1291:.
1281:.
1271:12
1269:.
1265:.
1249:^
1235:.
1225:.
1192:18
1190:.
1167:.
1159:.
1149:40
1147:.
1124:.
1116:.
1106:40
1104:.
1090:^
1076:.
1068:.
1056:56
1054:.
1050:.
1027:.
1017:19
1015:.
995:^
981:.
971:.
959:.
955:.
941:^
927:.
919:.
909:39
907:.
893:^
879:.
871:.
863:.
853:34
851:.
828:.
816:25
814:.
810:.
787:.
775:60
773:.
769:.
746:.
736:.
728:.
716:.
712:.
689:.
679:.
667:.
663:.
640:.
630:.
618:.
614:.
591:.
581:.
571:20
569:.
546:.
536:.
524:.
208:,
136:.
1994:.
1974::
1966::
1943:.
1931::
1905:.
1881::
1854:.
1822::
1803:.
1781::
1754:.
1734::
1726::
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1676::
1668::
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1619::
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1570::
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1531::
1502:.
1480::
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1399:.
1385::
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1338::
1330::
1307:.
1285::
1277::
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1202:.
1198::
1175:.
1163::
1155::
1132:.
1120::
1112::
1084:.
1062::
1035:.
1023::
989:.
967::
935:.
923::
915::
887:.
859::
836:.
822::
795:.
781::
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732::
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718:9
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675::
648:.
626::
599:.
585::
577::
554:.
532::
65:)
59:(
54:)
50:(
36:.
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