Patrícia Beldade
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Evolution and Development of Butterfly Wing Patterns
- the eyespots of Bicyclus anynana


During my Ph.D. work at the University of Leiden (The Netherlands), Paul Brakefield and myself used Bicyclus anynana to explore different (populational and organismal, evolutionary and developmental) processes that shape variation in morphology.

Using artificial selection on a large laboratory population, we explored the possibilities for changes in dorsal forewing eyespot size. Despite the evidence that all eyespots are genetically and developmentally coupled, artificial selection based on a single lab population produced changes in the two target eyespots in opposite directions and matched variation found across different Bicyclus species (pub#2). These results revealed much flexibility for independent changes in individual eyespots, probably resulting from the balance between the origin of all eyespots as a single developmental module and a history of selection favouring eyespot individuality (pub#4). Despite such flexibility, further illustrated by the phenotypes produced by different spontaneous mutations of large effect, there are obvious genetic correlations across eyespots on the same butterfly (pub#5).

The phenotypically divergent lines derived by selection were used in a number of experiments aiming at understanding the mechanisms underlying the production and maintenance of variation in eyespot size. These included laboratory mate choice experiments testing whether female B. anynana showed preference for males with different eyespot sizes, and manipulative experiments on pupae analyzing the changes in the cellular interactions underlying eyespot formation (pub#11).

The selection lines were also used to study the molecular genetic basis of variation in B. anynana eyespot size. This work was done in collaboration with Tony Long at the University of California at Irvine (CA, USA). Developmental biologists identified a number of genetic pathways involved in eyespot formation but little was known about which genes (if any) within the implicated pathways contributed to variation in eyespot patterns. We focused on the transcription factor Distal-less and showed that different alleles at this locus co-segregate with eyespot size in laboratory crosses between butterflies from different artificial selection lines (pub#1).
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Table of contents
Overview chapter
Samenvatting (NL)
Sumário (PT)

stellingen
dedicatória (PT)

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