title: In silico evo-devo: reconstructing stages in the evolution of animal segmentation.
abstract: BACKGROUND
  The evolution of animal segmentation is a major research focus within the field of evolutionary-developmental biology. Most studied segmented animals generate their segments in a repetitive, anterior-to-posterior fashion coordinated with the extension of the body axis from a posterior growth zone. In the current study we ask which selection pressures and ordering of evolutionary events may have contributed to the evolution of this specific segmentation mode.
  
  
  RESULTS
  To answer this question we extend a previous in silico simulation model of the evolution of segmentation by allowing the tissue growth pattern to freely evolve. We then determine the likelihood of evolving oscillatory sequential segmentation combined with posterior growth under various conditions, such as the presence or absence of a posterior morphogen gradient or selection for determinate growth. We find that posterior growth with sequential segmentation is the predominant outcome of our simulations only if a posterior morphogen gradient is assumed to have already evolved and selection for determinate growth occurs secondarily. Otherwise, an alternative segmentation mechanism dominates, in which divisions occur in large bursts through the entire tissue and all segments are created simultaneously.
  
  
  CONCLUSIONS
  Our study suggests that the ancestry of a posterior signalling centre has played an important role in the evolution of sequential segmentation. In addition, it suggests that determinate growth evolved secondarily, after the evolution of posterior growth. More generally, we demonstrate the potential of evo-devo simulation models that allow us to vary conditions as well as the onset of selection pressures to infer a likely order of evolutionary innovations.
authors.affiliation: Theoretical Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands.
authors.forename: Renske M A
authors.identifier: 0000-0002-1353-797X
authors.initials: RM
authors.lastname: Vroomans
authors.affiliation: Theoretical Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands.
authors.forename: Paulien
authors.initials: P
authors.lastname: Hogeweg
authors.affiliation: Theoretical Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands.
authors.forename: Kirsten H W J
authors.initials: KH
authors.lastname: Ten Tusscher
keywords: Axis extension; Bilaterian evolution; Determinate growth; In silico evolution; Posterior signalling; Segmentation