title: Drosophila wing melanin patterns form by vein-dependent elaboration of enzymatic prepatterns.
abstract: BACKGROUND
  Animal melanin patterns are involved in diverse aspects of their ecology, from thermoregulation to mimicry. Many theoretical models have simulated pigment patterning, but little is known about the developmental mechanisms of color pattern formation. In Drosophila melanogaster, several genes are known to be necessary for cuticular melanization, but the involvement of these genes in melanin pattern evolution is unknown. We have taken a genetic approach to elucidate the developmental mechanisms underlying melanin pattern formation in various drosophilids.
  
  
  RESULTS
  We show that, in D. melanogaster, tyrosine hydroxylase (TH) and dopa decarboxylase (DDC) are required for melanin synthesis. Ectopic expression of TH, but not DDC, alone was sufficient to cause ectopic melanin patterns in the wing. Thus, changes in the level of expression of a single gene can result in a new level of melanization. The ontogeny of this ectopic melanization resembled that found in Drosophila species bearing wing melanin patterns and in D. melanogaster ebony mutants. Importantly, we discovered that in D. melanogaster and three other Drosophila species these wing melanin patterns are dependent upon and shaped by the circulation patterns of hemolymph in the wing veins.
  
  
  CONCLUSIONS
  Complex wing melanin patterns are determined by two distinct developmental mechanisms. Spatial prepatterns of enzymatic activity are established late in wing development. Then, in newly eclosed adults, melanin precursors gradually diffuse out from wing veins and are oxidized into dark brown or black melanin. Both the prepatterning and hemolymph-supplied components of this system can change during evolution to produce color pattern diversity.
authors.affiliation: Laboratory of Molecular Biology, Howard Hughes Medical Institute, University of Wisconsin, Madison 53706, USA.
authors.forename: J R
authors.initials: JR
authors.lastname: True
authors.forename: K A
authors.initials: KA
authors.lastname: Edwards
authors.forename: D
authors.initials: D
authors.lastname: Yamamoto
authors.forename: S B
authors.initials: SB
authors.lastname: Carroll
mesh_terms: D000818:Animals; D005075:Biological Evolution; D019521:Body Patterning / Q000187:drug effects; D004268:DNA-Binding Proteins / Q000235:genetics; D004296:Dopa Decarboxylase / Q000096:biosynthesis / Q000235:genetics / Q000502:physiology; D004298:Dopamine / Q000494:pharmacology; D004330:Drosophila / Q000033:anatomy & histology / Q000201:enzymology / Q000378:metabolism*; D029721:Drosophila Proteins; D005260:Female; D015870:Gene Expression; D019476:Insect Proteins / Q000235:genetics; D008297:Male; D008543:Melanins / Q000096:biosynthesis*; D009154:Mutation; D010641:Phenotype; D013045:Species Specificity; D014446:Tyrosine 3-Monooxygenase / Q000096:biosynthesis / Q000235:genetics / Q000502:physiology; D014680:Veins / Q000033:anatomy & histology / Q000378:metabolism; D014921:Wings, Animal / Q000098:blood supply* / Q000201:enzymology / Q000378:metabolism*
chemical_list: D004268:DNA-Binding Proteins; D029721:Drosophila Proteins; D019476:Insect Proteins; D008543:Melanins; C116191:e protein, Drosophila; D014446:Tyrosine 3-Monooxygenase; D004296:Dopa Decarboxylase; D004298:Dopamine