title: Mealybugs nested endosymbiosis: going into the 'matryoshka' system in Planococcus citri in depth.
abstract: BACKGROUND
  In all branches of life there are plenty of symbiotic associations. Insects are particularly well suited to establishing intracellular symbiosis with bacteria, providing them with metabolic capabilities they lack. Essential primary endosymbionts can coexist with facultative secondary symbionts which can, eventually, establish metabolic complementation with the primary endosymbiont, becoming a co-primary. Usually, both endosymbionts maintain their cellular identity. An exception is the endosymbiosis found in mealybugs of the subfamily Pseudoccinae, such as Planococcus citri, with Moranella endobia located inside Tremblaya princeps.
  
  
  RESULTS
  We report the genome sequencing of M. endobia str. PCVAL and the comparative genomic analyses of the genomes of strains PCVAL and PCIT of both consortium partners. A comprehensive analysis of their functional capabilities and interactions reveals their functional coupling, with many cases of metabolic and informational complementation. Using comparative genomics, we confirm that both genomes have undergone a reductive evolution, although with some unusual genomic features as a consequence of coevolving in an exceptional compartmentalized organization.
  
  
  CONCLUSIONS
  M. endobia seems to be responsible for the biosynthesis of most cellular components and energy provision, and controls most informational processes for the consortium, while T. princeps appears to be a mere factory for amino acid synthesis, and translating proteins, using the precursors provided by M. endobia. In this scenario, we propose that both entities should be considered part of a composite organism whose compartmentalized scheme (somehow) resembles a eukaryotic cell.
authors.affiliation: Institut Cavanilles de Biodiversitat i Biologia Evolutiva, Universitat de València, Apartado Postal 22085, València 46071, Spain.
authors.forename: Sergio
authors.initials: S
authors.lastname: López-Madrigal
authors.forename: Amparo
authors.initials: A
authors.lastname: Latorre
authors.forename: Manuel
authors.initials: M
authors.lastname: Porcar
authors.forename: Andrés
authors.initials: A
authors.lastname: Moya
authors.forename: Rosario
authors.initials: R
authors.lastname: Gil
mesh_terms: D000818:Animals; D020562:Betaproteobacteria / Q000302:isolation & purification / Q000502:physiology*; D004269:DNA, Bacterial / Q000737:chemistry / Q000235:genetics; D004755:Enterobacteriaceae / Q000302:isolation & purification / Q000502:physiology*; D016680:Genome, Bacterial; D006430:Hemiptera / Q000382:microbiology* / Q000502:physiology*; D008969:Molecular Sequence Data; D017422:Sequence Analysis, DNA; D013559:Symbiosis
chemical_list: D004269:DNA, Bacterial
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