title: Beyond genomic variation--comparison and functional annotation of three Brassica rapa genomes: a turnip, a rapid cycling and a Chinese cabbage.
abstract: BACKGROUND
  Brassica rapa is an economically important crop species. During its long breeding history, a large number of morphotypes have been generated, including leafy vegetables such as Chinese cabbage and pakchoi, turnip tuber crops and oil crops.
  
  
  RESULTS
  To investigate the genetic variation underlying this morphological variation, we re-sequenced, assembled and annotated the genomes of two B. rapa subspecies, turnip crops (turnip) and a rapid cycling. We then analysed the two resulting genomes together with the Chinese cabbage Chiifu reference genome to obtain an impression of the B. rapa pan-genome. The number of genes with protein-coding changes between the three genotypes was lower than that among different accessions of Arabidopsis thaliana, which can be explained by the smaller effective population size of B. rapa due to its domestication. Based on orthology to a number of non-brassica species, we estimated the date of divergence among the three B. rapa morphotypes at approximately 250,000 YA, far predating Brassica domestication (5,000-10,000 YA).
  
  
  CONCLUSIONS
  By analysing genes unique to turnip we found evidence for copy number differences in peroxidases, pointing to a role for the phenylpropanoid biosynthesis pathway in the generation of morphological variation. The estimated date of divergence among three B. rapa morphotypes implies that prior to domestication there was already considerably divergence among B. rapa genotypes. Our study thus provides two new B. rapa reference genomes, delivers a set of computer tools to analyse the resulting pan-genome and uses these to shed light on genetic drivers behind the rich morphological variation found in B. rapa.
authors.forename: Ke
authors.initials: K
authors.lastname: Lin
authors.forename: Ningwen
authors.initials: N
authors.lastname: Zhang
authors.forename: Edouard I
authors.initials: EI
authors.lastname: Severing
authors.forename: Harm
authors.initials: H
authors.lastname: Nijveen
authors.forename: Feng
authors.initials: F
authors.lastname: Cheng
authors.forename: Richard G F
authors.initials: RG
authors.lastname: Visser
authors.forename: Xiaowu
authors.initials: X
authors.lastname: Wang
authors.forename: Dick
authors.initials: D
authors.lastname: de Ridder
authors.affiliation: Laboratory of Plant Breeding, Wageningen University, Droevendaalsesteeg 1, 6708 PB Wageningen, the Netherlands. guusje.bonnema@wur.nl.
authors.forename: Guusje
authors.initials: G
authors.lastname: Bonnema
mesh_terms: D001937:Brassica / Q000235:genetics*; D032461:Chromosomes, Plant; D019295:Computational Biology; D019143:Evolution, Molecular; D056726:Genetic Association Studies; D005819:Genetic Markers; D014644:Genetic Variation; D018745:Genome, Plant; D023281:Genomics / Q000379:methods; D005838:Genotype; D058977:Molecular Sequence Annotation; D005810:Multigene Family; D010641:Phenotype
chemical_list: D005819:Genetic Markers