title: Adaptive developmental plasticity: compartmentalized responses to environmental cues and to corresponding internal signals provide phenotypic flexibility.
abstract: BACKGROUND
  The environmental regulation of development can result in the production of distinct phenotypes from the same genotype and provide the means for organisms to cope with environmental heterogeneity. The effect of the environment on developmental outcomes is typically mediated by hormonal signals which convey information about external cues to the developing tissues. While such plasticity is a wide-spread property of development, not all developing tissues are equally plastic. To understand how organisms integrate environmental input into coherent adult phenotypes, we must know how different body parts respond, independently or in concert, to external cues and to the corresponding internal signals.
  
  
  RESULTS
  We quantified the effect of temperature and ecdysone hormone manipulations on post-growth tissue patterning in an experimental model of adaptive developmental plasticity, the butterfly Bicyclus anynana. Following a suite of traits evolving by natural or sexual selection, we found that different groups of cells within the same tissue have sensitivities and patterns of response that are surprisingly distinct for the external environmental cue and for the internal hormonal signal. All but those wing traits presumably involved in mate choice responded to developmental temperature and, of those, all but the wing traits not exposed to predators responded to hormone manipulations. On the other hand, while patterns of significant response to temperature contrasted traits on autonomously-developing wings, significant response to hormone manipulations contrasted neighboring groups of cells with distinct color fates. We also showed that the spatial compartmentalization of these responses cannot be explained by the spatial or temporal compartmentalization of the hormone receptor protein.
  
  
  CONCLUSIONS
  Our results unravel the integration of different aspects of the adult phenotype into developmental and functional units which both reflect and impact evolutionary change. Importantly, our findings underscore the complexity of the interactions between environment and physiology in shaping the development of different body parts.
authors.affiliation: Instituto Gulbenkian de Ciência, Rua da Quinta Grande 6, P-2780, Oeiras, Portugal. armateus@igc.gulbenkian.pt.|Institute of Biology, Leiden University, Sylviusweg 72, 2333, BE Leiden, The Netherlands. armateus@igc.gulbenkian.pt.
authors.forename: Ana Rita A
authors.initials: AR
authors.lastname: Mateus
authors.affiliation: Instituto Gulbenkian de Ciência, Rua da Quinta Grande 6, P-2780, Oeiras, Portugal. mmpita@igc.gulbenkian.pt.|School of Informatics and Computing, Indiana University, 919 East Tenth Street, Bloomington, IN, 47408, USA. mmpita@igc.gulbenkian.pt.
authors.forename: Manuel
authors.initials: M
authors.lastname: Marques-Pita
authors.affiliation: Institute of Biology, Leiden University, Sylviusweg 72, 2333, BE Leiden, The Netherlands. vicencio-oostra@dds.nl.|Department of Zoology, Cambridge University, Downing Street, Cambridge, CB2 3EJ, UK. vicencio-oostra@dds.nl.
authors.forename: Vicencio
authors.initials: V
authors.lastname: Oostra
authors.affiliation: Instituto Gulbenkian de Ciência, Rua da Quinta Grande 6, P-2780, Oeiras, Portugal. elafuente@igc.gulbenkian.pt.
authors.forename: Elvira
authors.initials: E
authors.lastname: Lafuente
authors.affiliation: Institute of Biology, Leiden University, Sylviusweg 72, 2333, BE Leiden, The Netherlands. pb499@cam.ac.uk.|Department of Zoology, Cambridge University, Downing Street, Cambridge, CB2 3EJ, UK. pb499@cam.ac.uk.
authors.forename: Paul M
authors.initials: PM
authors.lastname: Brakefield
authors.affiliation: Laboratory of Genetics, Wageningen University, Droevendaalsesteeg 1, 6708, PB Wageningen, The Netherlands. bas.zwaan@wur.nl.
authors.forename: Bas J
authors.initials: BJ
authors.lastname: Zwaan
authors.affiliation: Instituto Gulbenkian de Ciência, Rua da Quinta Grande 6, P-2780, Oeiras, Portugal. pbeldade@igc.gulbenkian.pt.|Institute of Biology, Leiden University, Sylviusweg 72, 2333, BE Leiden, The Netherlands. pbeldade@igc.gulbenkian.pt.
authors.forename: Patrícia
authors.initials: P
authors.lastname: Beldade
mesh_terms: D000222:Adaptation, Physiological / Q000235:genetics*; D000818:Animals; D002080:Butterflies / Q000235:genetics / Q000502:physiology*; D003463:Cues; D004777:Environment; D019143:Evolution, Molecular; D005838:Genotype; D006728:Hormones / Q000502:physiology; D010641:Phenotype; D014921:Wings, Animal / Q000502:physiology
chemical_list: D006728:Hormones