title: Recurrent evolution of heat-responsiveness in Brassicaceae COPIA elements.
abstract: The mobilization of transposable elements (TEs) is suppressed by host genome defense mechanisms. Recent studies showed that the cis-regulatory region of Arabidopsis thaliana COPIA78/ONSEN retrotransposons contains heat-responsive elements (HREs), which cause their activation during heat stress. However, it remains unknown whether this is a common and potentially conserved trait and how it has evolved.
  
  
  
  We show that ONSEN, COPIA37, TERESTRA, and ROMANIAT5 are the major families of heat-responsive TEs in A. lyrata and A. thaliana. Heat-responsiveness of COPIA families is correlated with the presence of putative high affinity heat shock factor binding HREs within their long terminal repeats in seven Brassicaceae species. The strong HRE of ONSEN is conserved over millions of years and has evolved by duplication of a proto-HRE sequence, which was already present early in the evolution of the Brassicaceae. However, HREs of most families are species-specific, and in Boechera stricta, the ONSEN HRE accumulated mutations and lost heat-responsiveness.
  
  
  
  Gain of HREs does not always provide an ultimate selective advantage for TEs, but may increase the probability of their long-term survival during the co-evolution of hosts and genomic parasites.
authors.affiliation: Department of Plant Breeding and Genetics, Max Planck Institute for Plant Breeding Research, Cologne, 50829, Germany.|Present address: Department of Plant Physiology, Ruhr-University Bochum, Bochum, Germany.
authors.forename: Björn
authors.initials: B
authors.lastname: Pietzenuk
authors.affiliation: Department of Plant Breeding and Genetics, Max Planck Institute for Plant Breeding Research, Cologne, 50829, Germany.|Department of Crop Science, Federal University of Rio Grande do Sul, Porto Alegre, RS, 91540000, Brazil.
authors.forename: Catarine
authors.initials: C
authors.lastname: Markus
authors.affiliation: Department of Plant Biology, University of Geneva, Sciences III, 30 Quai Ernest-Ansermet, 1211, Geneva 4, Switzerland.|Present address: The Sainsbury Laboratory, University of Cambridge, Cambridge, UK.
authors.forename: Hervé
authors.initials: H
authors.lastname: Gaubert
authors.affiliation: Department of Plant Breeding and Genetics, Max Planck Institute for Plant Breeding Research, Cologne, 50829, Germany.|Present address: Cardiovascular proteomics, Centro Nacional de Investigaciones Cardiovasculares, Madrid, 28029, Spain.
authors.forename: Navratan
authors.initials: N
authors.lastname: Bagwan
authors.affiliation: Department of Crop Science, Federal University of Rio Grande do Sul, Porto Alegre, RS, 91540000, Brazil.
authors.forename: Aldo
authors.initials: A
authors.lastname: Merotto
authors.affiliation: UMR1345 IRHS, Université d'Angers, INRA, Université Bretagne Loire, SFR4207 QUASAV, 49045, Angers, France.
authors.forename: Etienne
authors.initials: E
authors.lastname: Bucher
authors.affiliation: Department of Plant Breeding and Genetics, Max Planck Institute for Plant Breeding Research, Cologne, 50829, Germany. pecinka@mpipz.mpg.de.
authors.forename: Ales
authors.initials: A
authors.lastname: Pecinka
mesh_terms: D017360:Arabidopsis / Q000235:genetics / Q000502:physiology; D019607:Brassicaceae / Q000235:genetics*; D019143:Evolution, Molecular; D018506:Gene Expression Regulation, Plant; D018745:Genome, Plant; D018869:Heat-Shock Response / Q000235:genetics; D006358:Hot Temperature; D010802:Phylogeny; D018626:Retroelements / Q000235:genetics*; D020079:Terminal Repeat Sequences / Q000235:genetics*
chemical_list: D018626:Retroelements
keywords: Brassicaceae; COPIA; Evolution; Heat stress; ONSEN